Skip to main content

Full text of "Bulletin of the British Museum (Natural History) Zoo Supp"

See other formats


) IS 












24 June 1955 



23 August 1955 



24 June 1955 



19 August 1955 



10 January 1956 



17 February 1956 



21 February 1956 



24 February 1956 



21 February 1956 



27 March 1956 










No. i. A review of the Notostraca. By alan r. longhurst i 

No. 2. The Polychaete fauna of the Gold Coast. By norman tebble 59 

No. 3. A revision of the Octopodinae in the collections of the British Museum. 


No. 4. A revision of the family of Epicriidae (Acarina-Mesostigmata) . By 

G. OWEN EVANS. (Pis. 1-2) 169 

No. 5. The Monk seals (Genus Monachus) . By judith e. king. (Pis. 3-8) 201 

No. 6. The Cephalopoda of Madeira. Records and distribution. By 

w. j. rees and g. e. maul 257 

Notes on the European species of Eledone with especial reference 

to eggs and larvae. By w. j. rees. (Pis. 9-10) 283 

No. 7. The monotypic genera of Cichlid fishes in Lake Victoria. By 


No. 8. A revision of the Hydroid genus Perigonimus M. Sars, 1846. By 

w. j. rees 335 

On three northern species of Hydr actinia. By w. j. rees. (Pis. 
n-12) 351 

No. 9. The lizard genus Aprasia ; its taxonomy and temperature-correlated 

variation. By h. w. parker 363 

No. 10. Birds collected by Mr. F. Shaw-Mayer in the Central Highlands of 

New Guinea 1950-1951. By r. w. sims. (Pis. 13-14) 387 

Index to Volume 3 439 

P. 4,6. The field note under Loria loria amethyslina relates to CmmopMlus mac S re g orii sanguineus 
on p. 427. 


2 JUL 1955 





ZOOLOGY Vol. 3 No. i 

LONDON: 1955 




(Bedford College, University of London) 

Pp. 1-57 ; 16 Text-figures 

ZOOLOGY Vol. 3 No. 1 

LONDON: 1955 

(NATURAL HISTORY), instituted in 1949, is 
issued in five series, corresponding to the Departments 
of the Museum, and an Historical Series. 

Parts will appear at irregular intervals as they 
become ready. Volumes will contain about three or 
four hundred pages, and will not necessarily be com- 
pleted within one calendar year. 

This paper is Vol. 3, No. 1 of the Zoological Series. 


Issued June, 1955 Price Twelve Shillings and Sixpence 



Materials . 
Systematic Characters 


Total Size 
Colour . 


(5) Carapace armature 

(6) Supra-antennal crest 

(7) Eyes, dorsal organ 

(8) Telson . 

(9) Segmental armature 

(10) Appendages 

(11) Furca . 
Protein Specificity 
Cytology . 








This paper reviews the known species of Notostraca on the basis of a large collection of museum 
material and on information from the literature. The genus Triops is reduced to four, and the 
genus Lepidurus to rive species. 

The species of Triops are separated on the armature of the telson and on the presence or 
absence of a second maxilla ; in each species there is considerable variation in the segment 
number, and in the many structures whose variation is shown to be correlated with this number. 

The segment number is more constant in Lepidurus and fewer structures are correlated with 
its variation ; this genus appears to fall into two groups, as Linder (1952) suggested, on the 
basis of the segment number. A new species of Lepidurus is described from Russia. 

In addition to the analysis of the validity of the systematic characters from work on preserved 
specimens, evidence derived from the growth and development of living animals, their cytology, 
their reproduction and their protein specificity are also considered. 


The Notostraca are an order of Euphyllopod Crustacea with shield-shaped carapace, 
consisting of a single family with two genera; Triops (= Apus) and Lepidurus. 
They usually occur in temporary pools of fresh or brackish water, being most common 
in the drier parts of the world where the surface water is often of a temporary nature. 
Their adaptation to a temporary habitat has enabled their drought-resistant eggs 
to become efficient agents of passive dispersal, so that populations occur on remote 
oceanic islands and are apparently found wherever there are suitable pools. 

ZOOL. 3, 1. I 


These animals are of interest to the general zoologist not only on account of their 
primitive nature, but also because they exemplify an animal in evolutionary stagna- 
tion. The family has been in existence since the Permian, and forms from the 
Triassic are almost indistinguishable from one of the extant species. 

Individually Notostraca are notoriously variable and differences in the armature 
of spines on the exoskeleton, or in bodily proportions, can be found in any pair of 
animals, even those from the same pool. This, together with the lack of conspicuous 
morphological discontinuities within the genera, makes the group a " difficult " one 
systematically, and has resulted in the excessive number of descriptions — of 
specimens rather than species — with which the synonymies are now burdened. 

There have been several revisions of the group as a whole, but none later than that 
of Simon (1886) who revised the species then known to him. Barnard (1929), Linder 
(1952) and Tiwari (1952) have reviewed regional material from South Africa, North 
America and India respectively, and have tended towards a reduction in the number 
of species recognized, placing many names into the synonymies. This trend has 
been general in recent years but has by no means prevented the description of new 
species on inadequate grounds. 

Because of their interest from a systematic and evolutionary point of view, their 
geological age, their passive distribution, and their restricted habitat, it was evident 
that a review of the Notostraca based on a comprehensive collection of preserved 
specimens and on a study of the growth and cytology of living animals was desirable ; 
one of the points of agreement between recent authors on the group has been the 
need for such a revision. 


The preserved specimens of Notostraca on which this revision is based consist of 
more than 200 samples, of which about 160 are of Triops ; the number of specimens 
in each sample varies and the majority contain less than 10 individuals, although 
a few have more than 100 specimens from the same locality. 

The bulk of this material is formed by the collections of the British Museum and 
of the Museum Nationale d'Histoire Naturelle, Paris ; the remainder was on loan 
from the U.S. National Museum in Washington, the Western Australian Museum 
in Perth, the Zoological Survey of India in Calcutta, and the Museo de Ciencias 
Naturalis in Madrid. A few samples of preserved Notostraca were sent to me by 
private individuals. 

The living material was reared in the laboratory from eggs in dried mud sent to 
me from phyllopod pools all over the world. Many contained no viable eggs, but 
successful hatchings were obtained from the following samples (collector's name in 
parentheses) : Triops cancriformis — Sweden (P. Ardo), England (A. R. Longhurst), 
France (D. Schachter), Italy (H. M. Fox) ; T. granarius — Johannesburg (van der 
Horst), Grahamstown (J. Omer-Cooper) , Iraq (A. James) ; T. longicaudatus — 
California (L. E. Rosenberg) ; T. australiensis — Ayers Rock (I. Thomas, F. McNeill, 
A. L. Rose), Kalgoorlie, Ballodonia (A. R. Main) ; Lepidurus arcticus — Iceland 
(H. Moore) ; L. apus — New Zealand (A. Lysaght, E. Percival). 

Living cultures of Notostraca were successfully maintained in the laboratory 


under reasonably standard conditions. Larvae hatched readily from the mud 
samples when these were placed in a tank of clean water (either filtered pond water 
or dechlorinated tap water) which were maintained at about 20° C, air being bubbled 
through the water. If an excess of mud was put in a tank bacterial growth was very 
rapid and the culture was soon lost ; about J in. on the bottom of a 24 in. by 12 in. 
tank proved suitable. 

The larvae fed at first on the organic content of the mud and if the production of 
diatoms and Protozoa was poor in the culture in its early stages an addition of 
cultured Chlorella was very beneficial to the growth of the larvae. From about 1 cm. 
in length growth was more rapid if additional food was provided ; this consisted of 
chopped annelid worms (Tubifex), live Daphnia and an artificial food. This last was 
made up from equal parts by weight of dried Daphnia, grass-flour, and Bemax (a 
proprietary cereal food) ground together into a fine powder and suspended in calcium 
alginate jelly. This was then chopped and formed a satisfactory basic diet for adult 

The eggs laid in the cultures were hatched by allowing the mud which contained 
them to dry out slowly ; on re-wetting, a high proportion of them hatched success- 
fully. A more rapid and effective way of maintaining a culture was to collect the 
eggs as they were laid and to transfer them to a beaker of clean tap water with an 
addition of 30-50% of glass-distilled water ; this low osmotic pressure medium 
induced hatching after 12-14 days without prior drying, in the same manner as 
Hall (1953) has described for the eggs of Chirocephalus diaphanus. The larvae live 
for only a few moments in this water however, and must be removed from it at once. 

I am most grateful to those who sent me samples of mud (including those from 
which I was unable to hatch Notostraca and which are not listed above) and to 
those people who collected and sent preserved specimens. My thanks are also due to 
the Authorities of the Museums listed above who lent me material, and especially 
to the Trustees and Staff of the British Museum (Natural History) and the Museum 
Nationale d'Histoire Naturelle, who generously gave me facilities for examining 
their collections. I am particularly indebted to Dr. J. P. Harding of the British 
Museum who has given me invaluable help during the whole time that I have been 
studying the group. 

The work on living animals was done in the Zoology Department of Bedford 
College, University of London under the supervision of Professor H. Munro Fox, to 
whom I am grateful for much assistance. 

A maintenance grant from the Department of Scientific and Industrial Research, 
and a special grant from the Central Research Fund of the University of London 
were received during the work. 

Abbreviations. — In the text the museums listed above are referred to by the follow- 
ing sets of initials : 

BMNH . . . London. 




. Paris. 


. Perth. 

ZSI . 

. Calcutta. 



The systematics of the Notostraca have been based almost entirely on characters 
of the setae and spines which comprise the armature of the exoskeleton, and on the 
proportions of various parts of the body — the endites, the furca and the carapace. 

The validity of these characters was discussed in Barnard's revision of the South 
African forms (Barnard, 1929). More recently, the review by Linder (1952) further 
explored this field and has added several characters which prove useful in separating 
species. The description of new species from European and North African material 
by Ghigi (1921) initiated a discussion on the validity of the characters which he 
used, to which Colosi (1922), Gurney (1923), and Gauthier (1933, 1934) contributed. 

Before any character can properly be used in systematics it is essential that its 
variation in adult individuals be known, and that its development or change during 
the growth of a single individual should have been studied ; in the present review 
the validity of as many as possible of the systematic characters has been studied 
and several new ones are proposed. 

Previous authors have been concerned almost without exception with the variation 
of characters in samples of adult individuals, and it has been possible to extract 
much useful information about this from the literature, which has been combined 
with that obtained from the study of the museum material. 

A pure line of individuals from a hermaphrodite Triops cancriformis from Britain 
was raised under standard conditions ; a large batch of eggs was collected from the 
parent, hatched, and reared at first in a single large culture dish, later being trans- 
ferred to individual dishes, each with the same volume of water and mud ; the 
dishes standing in a room thermostatically controlled at 20 C (± i° C). A surplus 
of the artificial food was always available to the animals. From this culture 25 
individuals were preserved and their variation studied ; all these animals were 
preserved after growth changes had ceased in the characters to be studied. 

The change of several characters during growth was studied by Linder (1952) in 
museum material ; he based his conclusions on specimens which were about to 
ecdyse, in which both the old and the new exoskeletons could be observed. In the 
present review living material of all species of Triops was studied during growth 
from larva to adult ; the changes in relative proportions during growth have been 
examined and the size at which the exoskeletal armature becomes stable determined. 

A start has also been made on the effects of environmental factors on morphology, 
but the experiments have not been very successful. Triops is a difficult animal to 
grow under precisely standardized conditions, for it has not been possible to rear 
them without a little mud in their dishes even when they are fed artificially. Main 
(1953) has suggested that so-called specific differences in Australian forms may be 
due to differences in the salinity of the medium ; I have made attempts to grow 
Triops at salinities near those that Main found in the field, but the animals have 
rapidly died each time. The temperature at which the animals were reared might 
be expected to have some effect on their form, and so a pure line of T. cancriformis 
was grown at a temperature (25-28 C) near their lethal point, but these animals 
showed little difference from those grown at normal temperatures. 


Notostraca were grown in the laboratory under very diverse conditions of tempera- 
ture, food, and vessel size, but all have remained remarkably uniform in their 
morphology. Daphnia under parallel conditions would show very marked morpho- 
logical changes, and it is probably safe to assume that environmental factors can 
affect the morphology only after a very considerable number of generations — as 
salinity affects the form of Artemia salina only after several years (Schmankiewitsch, 


The various characters which have been used in systematics are now examined 

(1) Total size 

Samples of adult Notostraca usually contain individuals of very different sizes, 
and it is difficult to determine whether there is a normal adult size for any species. 
The growth rates of animals in a batch of Triops reared in the laboratory are also 
very variable (Fox, 1949) as the pure line of T. cancriformis showed very clearly ; 
after a few days growth they varied enormously in size ; 33 individuals were between 
2*0 and 3*5 mm. carapace length, while one was only 1*5 mm., and another 5-0 mm. 
Spandl (1926) gives growth curves for individual T. cancriformis, maintaining that 
those which grow the fastest become the largest, which is what one would expect 
if the growth rate depended on the level of nutrition. 

The usual adult size for all species seems to be between 15 mm. and 30 mm. in 
carapace length, the growth curve flattening out somewhere between these two 

The total size has occasionally been used as a systematic character. 

Bowkiewicz (1923) suggests that giant Triops which he saw in Siberia might be a 
new species ; large individuals of other species occur up to 40 mm in carapace length 
and probably correspond to the giant specimens of Anostraca, which Sellier and 
Morice (1946) have shown to be cytologically similar to normal individuals in one 
species (Chirocephalus diaphanus). 

It is difficult to find a suitable measure of the total size of individuals in order 
that relative proportions of endites, etc. may be compared during growth in different 
samples. The total length has been shown to be quite useless in preserved specimens 
(Barnard, 1929 ; Somme, 1934 ; Linder, 1952) and is difficult to measure in living 
animals, which have considerable powers of contraction. The measurements made 
on living animals also indicate that there is some increase in length during an instar. 
Somme (1934) showed that the median carapace length is more reliable for it is 
little affected by the action of the preservative. 

It has now been possible to show that the growth of the carapace is isometric and 
is therefore a valid measurement of size at all ages (p. n). 

Unfortunately, the ratio of carapace length /total length varies between samples 
and so this measurement is valid only in comparisons within a sample. 

Generally, Lepidurus spp. tend to be smaller than Triops spp. and Lepidurus 
arcticus is usually smaller than the rest, but probably more rarely achieves its 
maximum size than the non-boreal species. 


(2) Colour 

Living Notostraca are frequently brightly coloured, the colours being due to two 
main pigments within the body and the brown colour of the exoskeleton. 

The internal pigments are haemoglobin, which is present in solution in the blood 
(Regnard and Blanchard, 1883), and a dark blue-green pigment which occurs in 
connective tissue in many parts of the body ; this pigment is of unknown composi- 
tion, but is similar in nature to that which occurs in some Ostracods (Fox, 1955). 

The concentration of haemoglobin varies inversely with the oxygen tension of the 
medium in which the animal is living (Fox, 1949), and large animals in poorly aerated 
conditions may have so dense a concentration that the animal appears to be deep 
red in colour. 

The blue-green pigment appears to be more abundant in Lepidurus than in Triops 
and the animal is sometimes deep green in colour (hence Lepidurus viridis, 
Baird). In Triops it may be completely absent, though a mid-dorsal patch on the 
carapace and smaller areas on the bases of the thoracic appendages are generally 
present. It seems to be more abundant in animals which have grown rapidly and 
under good conditions, when it forms a dark marbling on the carapace and is well 
distributed elsewhere. The only living specimens of T. australiensis that I have 
seen have been without this pigment, but this is unlikely to have any significance. 

The newly hatched larvae of both genera may be densely coloured with a carotenoid 
pigment, as Fox (1949) reported for Triops cancriformis ; in successive generations 
of the same stock the colour of the larvae is very variable and probably depends on 
the feeding of the parents ; well fed females usually contain eggs which are pink 
with carotenoid, while in starved specimens they are white. 

The egg shells of Triops are bright red in colour, for a red haemochromogen is 
present in them (Fox, 1955) , this pigment is secreted by the follicle ducts of the 
ovary, and is brightest in colour in newly laid eggs, becoming duller after they have 
been dried. 

A bright violet pigment occurs in the egg shell of Lepidurus arcticus (H. Moore, 
personal communication), while other species in this genus have eggs of the same 
colour as in Triops. 

(3) Body-length, segmentation, and number of appendages. 

The carapace of Notostraca is attached only to the head region, and the thorax 
and abdomen are completely free from it ; the post-carapace region is divided into 
a number of segments or body rings, which have considerable powers of telescoping. 
The first eleven segments, of which the first is incomplete dorsally, normally each 
bear one pair of appendages ventrally, and together comprise the thorax. 

The number of post-thoracic, or abdominal, segments is variable and the series 
of appendages is continued along them ; a few segments at the posterior end of the 
abdomen bear no appendages. 

linder (1952) has analysed a great deal of North American material and has to a 
great extent elucidated the relationships between the number of segments, the 
number of appendages and the number of apodous segments posteriorly. He con- 


eludes that the number of segments and the number of appendages are the results 
of two quite separate growth processes for a number of reasons : the production of 
segments and appendages in the larvae proceed at different rates ; the boundaries 
of the appendage bearing segments are not complete ventrally ; an aberration in 
which the segments are spiral instead of annular does not upset the arrangement of 
the appendages ; no correlation can be found between the number of appendages 
and the segments bearing them ; and the appendage series may end anywhere 
along the length of a segment. 

While studying the development of the systematic characters during growth I 
found that the larvae of Triops complete their segmentation by the 5th or 6th 
instar, but that the series of appendages continue to increase and encroach posteriorly 
on to fresh segments until the 8th or 9th instar, after which the number of apodous 
segments remains constant. After this the number of appendages may continue to 
increase for a few instars but come to occupy no further segments. So after the 9th 
instar the number of segments and the number which are apodous may be taken as 
fixed in an individual, so that these could validly be used as characters in animals 
of more than 3-4 mm. in median carapace length. 

The total number of segments varies in Triops from 32-44, and in Lepidurus from 
26-34 '> these results are based on the museum material combined with information 
from the literature. Throughout this paper the figures given do not include the 
telson which is considered to be post-segment al, and incomplete segments are 
included in the count. 

The figures for Triops show that the variation is continuous throughout its range, 
and separation into groups on the number of segments would be quite arbitrary ; 
the results for females from all sources illustrates this — 

No. of segments . 32 33 34 35 36 37 38 39 40 41 42 43 
No. of occurrences . 24 31 19 12 11 17 22 12 13 15 10 7 

The bottom row of numbers — the number of occurrences — is obtained by counting 
the number of samples in which each segment-number occurs ; this prevents undue 
weight being given to large samples, which are not evenly distributed along the range 
of variation. 

The apparent bimodality of these figures is due to the fact that one species — 
Triops cancriformis — has a range of segment variation which covers only a part of 
that of the other species. This species occurs in Europe and naturally predominates 
in the collections, thus increasing the number of occurrences at the lower end of 
the scale. T. cancriformis has a variation of 32-35, while the other species probably 
run from 32-43. 

By far the greatest number of Lepidurus examined fall within the range of 25-29 
segments ; I have seen only two samples with more, L. lynchi Linder and L. batesoni 
sp. n. This is in accordance with Linder's findings, for he suggested that there 
might be two groups of species within Lepidurus based on the number of segments. 

The variation within samples bears out the theory that the number of segments 
may be of use in the systematics of Lepidurus but not of Triops ; in the pure line 
of T. cancriformis there was a variation of 3 in this character (33-35), and variations 


of 2, 3, or 4 are usual in samples of preserved specimens. In Lepidurus, however, 
possibly because there are fewer segments, the variation is smaller, rarely being 
more than 2 in each sample, many samples showing none. 

The number of appendages confirms the above grouping of Lepidurus ; the shorter 
bodied group has from 35-48 appendages, the longer from 39-71 ; these figures are 
based on Linder's data with the addition of counts made on the material I have 
examined. But in Triops I can find no correlation between the number of legs and 
the number of segments, for high numbers of appendages occur in both short and 
long bodied specimens, and the converse is also true. In this genus I can find no 
significance in the numbers of appendages. 

The number of apodous segments is a secondary character depending on the 
interplay of the processes which control appendage and segment formation (Linder, 
1952). This is confirmed in my data, most commonly with males and females from 
the same sample ; the males tending to have a smaller number of appendages, the 
same number of segments and so a higher number of apodous segments than the 
females. Similarly, Lepidurus bilobatus has 33 segments, 60 pairs of appendages and 
6 apodous segments (Linder, 1952), while L. batesoni with the same number of 
segments, but only 39-52 appendages, has 8-9 apodous segments. 

The apodous segments were frequently counted in early descriptions of Notostracan 
species, and several specific distinctions have been based on small differences in this 
number. It is now known that there is so much variation in this character in Triops 
that it is useless as a systematic character ; the pure line T. cancriformis had a 
variation of three (5-7). It may be valid in some Lepidurus, and is of use in at least 
one specific distinction. 

A few general rules can now be drawn from the data on segmentation and appendage 

In both genera males often have a higher number of segments within a sample, 
while the reverse appears to be unknown. In 25 samples of Triops I found this to be 
the case, while in 21 the difference was insignificant. 

Certainly in Triops, and probably in Lepidurus, the males tend to have fewer 
appendages and so a higher number of apodous segments than the females ; in 44 
Triops samples the males had a higher number of apodous segments, in 7 there was 
no difference and in 1 the female had more. 

Specimens with high numbers of segments tend to have a high apodous number 
in both genera. 

Thus, it is obvious that these characters can be used in systematics only with a 
great deal of caution, and appear to be of more use in Lepidurus than in Triops. 

(4) Carapace 

The shape and size of the carapace in Notostraca varies considerably and the 
differences found have frequently been used in the past by S3 7 stematists to dis- 
tinguish species. Ghigi (1921) used the carapace shape as one of his arguments in 
separating Triops into two genera : Thriops and Proterothriops (sic) ; Barnard 
(1929) considered the shape to be a distinguishing feature between South African 
species of Triops, particularly between his Apus numidicus and A. namaquensis. On 


the other hand, Linder (1952) made no use of the carapace size and shape in his 
revision of the North American forms. The carapace is one of the structures which 
is usually described adequately in the earlier papers. 

In each individual the growth of the carapace from the earliest larva is probably 
isometric ; a number of individuals of Triops cancriformis and T. granarius were 
measured during the whole of their growth period and it was found that the ratio of 

5 IO 



Fig. i. Growth coefficient (k) of females of Triops cancriformis from Hampshire — five 

carapace length to total length remained the same throughout growth. In the 
former species the growth coefficient (k) of the carapace was 1/02, while in the latter 
species k = roi ; these results are shown graphically in Text-fig. 1. 

The basic variation in the carapace is its size relative to the total length of the 
animal ; in some forms it covers a much greater length of the body than in others ; 
correlated with this relative size-difference are other differences in shape and strength 


of the carapace ; in those specimens in which it is relatively small it is always more 
rounded in outline, flatter, and less strong than in those in which it is large. These 
differences hold good for both genera. 

The relationship between carapace length and total length depends both on 
changes in size of the carapace — the number of segments that it covers — and on 
changes in the number of abdominal segments. A comparison between two species 
of Triops made on the living animals illustrates this ; British female T. cancriformis 
have about 33 segments of which about 19 are exposed behind the carapace, which 
thus covers some 14 segments ; T. granarius from Johannesburg have, in the female, 
about 38 segments of which 27 are exposed, so that in this species the carapace 
covers about n segments. In the second species the increase in the number of 
exposed segments comes both from a shorter carapace and a larger number of 
abdominal segments. It is impossible to make such calculations on preserved 
specimens with any accuracy, because of contraction in the preservative. 

The carapace appears to cover about 11-14 segments in most specimens, for on 
these segments are borne the most anterior of the spines which occur on the margins 
of the segments in the exposed portions of the abdomen ; in all the specimens 
examined, the first of these spines appear on segments n-14, regardless of the total 
number of segments present. 

Within a species, males have smaller carapaces than females, although this 
dimorphism is less marked in Lepidurus than Triops, and in shorter bodied than in 
longer bodied specimens of the latter genus. This sexual dimorphism in the carapace 
has been noted by many authors. 

The dependence of the degree of this dimorphism on the number of segments 
means that while it is usually possible to distinguish males from females at a glance 
on this character within a population, males from short bodied populations may 
actually have longer carapaces than females from populations with high segment 
numbers. In the forms with the lowest numbers of segments the dimorphism may 
be so slight as to be virtually non-existent. 

The carapace is smaller, more round, and flatter in populations of both genera 
which have relatively high segment numbers ; this is especially well marked in 
Triops where the specimens with the highest numbers of segments have remarkably 
small carapaces (Text-fig. 2). 

Barnard (1929) maintained that the shape of the carapace was an absolute dif- 
ference between his Apus numidicus and A. namaquensis , which he found in the 
former species to be oval in shape and in the latter almost round ; he also gave data 
on the number of apodous segments which show that A. namaquensis is longer in 
the body than the other species, although some overlap occurred between the two. 
He was concerned only with South African material, but on examining specimens of 
these species — and of synonymous ones — from the whole of their range in Africa 
and Asia I find that there is no discontinuity in the variation of carapace-shape. 
The round carapaces and the oval are connected by populations of intermediate 
form (Text-fig. 2A-e). The differences that Barnard found were differences between 
the long and the short bodied forms of the same species. 

I can find no differences in the carapace shape or size which are not correlated 



Fig. 2 Correlation of form with number of segments, a-e, Triops granarius, progres- 
sive increase of segments (32-42) ; f, squared sulcus on some specimens of T. granarius 
from South Africa ; g-h, T. longicaudatus , few and many segments respectively ; 
1, Lepidurus apus ; j, L. batesoni. (3, 4, 5, — endites of first thoracic appendage, 
cp = carapace, t = telson, fc = furca, sap — supra-anal plate). 


with sex or the number of segments ; Triops cancriformis tends to have a lower 
number of segments than T. granarius and so its carapace tends to be longer, less 
flat, than in the other species ; and specimens of T. granarius with a low number of 
segments have carapaces similar in shape to those of T. cancriformis. Lepidurus spp. 
tend to have even shorter bodies and here the carapace may cover most of the 
abdomen, and may be so deep as to enclose the animal laterally. 

The shape of the sulcus, or posterior emargination, of the carapace has frequently 
been described and importance attached to it ; I find it to be very variable and can 
see no correlation with other characters ; it may be shallow and wide in both long 
or short bodied forms, or small and round in similar animals. A peculiar squared 
sulcus has been seen in several short bodied populations of Triops granarius from 
Africa (Text-fig. 2f), but this grades into more rounded forms and is obviously of 
no significance. Wide, shallow sulcus shapes occur most frequently in T. longicaudatus , 
but this is of very doubtful value in the systematics. 

The difficulty of an accurate classification of sulcus shapes makes it unlikely that 
this will ever be a useful character. 

(5) Carapace armature 

The carapace bears an armature of spines both scattered and localized. The whole 
outside surface may be smooth, or may bear scattered upright spines ; the dorsal 
carina frequently ends in a spine and may bear smaller spines along its length ; the 
sulcus generally bears a row of marginal spines, and the outside edge of the carapace 
may bear a similar row. 

The carapace armature shows a great deal of variation, and has been referred to 
very frequently in past descriptions ; some of this variation does seem to be of use 
systematically, but it must be used with caution and only as a confirmatory character 
in most cases. 

The armature as a whole varies in its strength and development even within a 
population, and in some animals the whole armature is more strongly developed 
than in others — in the former not only are the spines larger and stronger, but they 
are also more numerous. In Triops granarius, when the scattered surface spines are 
well developed, the sulcus spines are particularly strong, the carina is denticulate 
and the whole carapace is more rigid than in other specimens. In a population of T. 
cancriformis from Tunisia (MNHNP) the armature of the exoskeleton is extra- 
ordinarily weakly developed, though typical of this species in arrangement, and 
extreme examples in this sample have no carinal or sulcal spines at all — a most 
singular condition. 

The scattered spines on the surface of the carapace were features used by Sars in 
the description of two species of Triops (Apus trachyaspis and A. sculleyi, Sars, 
1899), but Barnard found that specimens bearing such spines occurred sporadically 
among South African material. I have found such specimens in samples of Triops 
granarius and T. longicaudatus, both as isolated individuals and as complete samples. 
Specimens in which this character is well developed are so conspicuous — the carapace 
having a prickly feel — that I shall refer to them as the trachy aspis-i orm of whichever 
species is involved. 


The terminal spine of the carina is most prominent in larval Lepidurus] I have seen 
specimens of L. arcticus, L. apus apus and L. apus viridis in which it is relatively 
enormous in the second and third instars (Text-fig. 13A). In larvae of Triops it 
develops later and is never as large as in young Lepidurus. The growth rate of this 
spine in Lepidurus must be strongly negatively allometric, for although it is present 
in most adults, it is relatively very much smaller in these than in the larvae ; in 
Triops, when it occurs, this spine has a positively allometric rate for it is quite small 
when it first appears, and becomes, in the adult, of a size relatively similar to that 
of adult Lepidurus. 

This terminal spine occurs in almost all adult Lepidurus and is absent in only a 
few. There appears to be no correlation between its absence and other characters ; 
L. batesoni and some specimens of L. apus apus and of L. apus lubbocki are without 
it, but these specimens of L. apus are otherwise quite normal. There is probably a 
variation in the growth rate of this spine to account for its disappearance in adults 
of a few populations. 

Fig. 3. Carapace of Lepidurus lynchi. a, typical form of carinal spines ; b, specimen 
with no carinal spines. 

In Triops there is a loose correlation between the occurrence of the terminal spine 
and other characters ; within a species it is more commonly present in short than 
in long bodied forms. There is some difference between species, too, and as might be 
expected it is almost always present in the relatively short bodied species, T. cancri- 
fortnis. It is present in the remaining species only in their short bodied forms. 

The carina may bear a series of spines, most commonly at the posterior end just 
anterior to the terminal spine. These have frequently been used in the past for 
systematics (Ghigi, 192 1 ; Linder, 1952) and they have proved to have value in 
certain cases. In Triops they may be present, or absent, in both long and short 
bodied forms, but there are some interesting differences within T. cancriformis. In 
this species a few small spines are generally present posteriorly, and these are more 
numerous and much stronger in specimens from Morocco and Southern Spain ; these 
are the populations referred to by Ghigi (192 1) as Thriops mauretanicus , but they 
are now considered to comprise a sub-species (sens. Mayr, et al) of the more wide- 
spread Triops cancriformis. Another sub-species of this species is characterized by 
the complete absence of carinal spines. 

In Triops australiensis and T. longicaudatus these spines are most frequently 


completely absent, but in a number of populations, that from the rice fields of Biggs 
County in California, for example, the whole carina bears an even row of very small 

Most specimens of Triops granarius are without carinal spines, and only a few of 
the short bodied specimens have an arrangement like that of T. cancriformis. Very 
rarely this species has a long row of very small denticles like those of T. longicaudatus. 

The majority of specimens of Lepidurus have perfectly smooth carinae but a 
remarkable series of large teeth along the carina occurs in L. lynchi, quite unlike 
anything else in the Notostraca (Text-fig. 3). 

The sulcus normally bears a marginal row of spines, the only exceptions being 
some of the specimens of Triops cancriformis from Tunisia mentioned earlier. There is 
a fairly clear correlation of the form of these spines with the number of segments in 
Triops, but not in Lepidurus ; they are larger and fewer in number in short bodied 
forms of the former genus. 

Their development in Triops is fairly clear ; the spines at the outside angles of 
the sulcus appear first, in the 4th or 5th instar, at which time the rest of the sulcus 
has a finely granulated margin. The first marginal spines appear in the 7th or 8th 
instar, and increase in number until about the 10th instar, after which time the 
number is fixed, though in a few specimens small subsequent additions may occur. 
There is little variation in this character in an individual after it is about 5*0 mm. 
in carapace length. 

Barnard (1929) considered that there was so much variation between individuals 
in the sulcus spines that their use in systematics was not justified, although earlier 
writers had placed much emphasis on them. Linder admitted this variation, but 
thought that in some cases specific differences could be found. 

I can discover no differences of the sulcal spines in Triops which can have any 
value in systematics, most of them are correlated merely with body length ; in T. 
granarius, short bodied forms have long, slightly curving spines and in longer bodied 
animals a larger number of small, blunt spines occurs. As in other characters T. 
cancriformis here resembles the shorter bodied T. granarius specimens. 

In Lepidurus, a correlation with body length was not observed ; some specimens 
with 28 and some with 33 segments had small squat spines, but the vast majority of 
samples of all body lengths have long spines similar to those of Triops cancriformis. 
An unusual arrangement occurs in Lepidurus apus packardi in which the margin 
is closely set with many small squat spines like those of larval Triops. 

The outer margin of the carapace in both genera normally bears a series of denticles 
which produce a finely serrated edge ; this is variable in development, and is normally 
stronger near the posterior angles of the carapace ; in only one case are these 
marginal denticles of any value systematically ; some specimens of Lepidurus 
lynchi bear a series of teeth along this edge which are very much larger than those 
of any other known form. 

The amount of individual variation which may be expected in the carapace 
armature of a population was illustrated by the pure line of Triops cancriformis ; 
here the number of posterior teeth on the carina varied from 2-10, the number of 
sulcus spines from 24-32, and all the specimens had a large terminal carina spine. 



(6) Supra-antennal crest 

On either side of the ventral surface of the head there is a ridge, behind which is 
set the first antenna ; this is the supra-antennal crest of Simon (1886), which Linder 
(1952) suggests may be worth study. I find it variable within a single species, in 
some specimens of Triops c. cancriformis it is denticulate and in others smooth and 
less prominent. It seems improbable that it is of importance systematically. 

(7) Eyes and dorsal organ 

In all adult Notostraca the dorsal surface of the head bears a pair of compound 
eyes, an ocellus, and the dorsal (or nuchal) organ. 


Fig. 4. Eyes and dorsal organs, a, Triops cancriformis ; b, T. australiensis 
Triops granarius ; d, Lepidurus batesoni ; B,F,L.apus; g, L. arcticus. 

c, c. 

The size and arrangement of these structures are given in most specific descriptions 
and considerable importance has been ascribed to them, in particular to the dorsal 
organ ; Barnard (1929) showed that the dorsal organ varied in South African 
material from a small round shape to a larger and triangular one. Further, he found 
that the small, round dorsal organs tended to be set on a tubercle in his animals, 
but that the triangular ones were less elevated and nearly flush with the surface of 
the head. These differences, to which he ascribed much importance in separating 
species, now appear to be another example of a character which is correlated with 
the number of segments, and which varies in a similar fashion in several species. 

In all the species of Triops the specimens with a low number of segments tend to 
have small, round and elevated dorsal organs, and as the number of segments 
increases so the dorsal organ becomes larger, less elevated, and more triangular in 
shape (Text-fig. 4C-c 2 ). 

This is well shown by Triops granarius ; in specimens from South Africa with only 
33 segments the dorsal organ is small and round, or slightly pear-shaped, as it is in 
zool. 3, 1. 2 


specimens with 36-37 segments from Chufoo, China (both BMNH) ; in longer 
bodied specimens it is much larger and more triangular. In T. cancriformis the usual 
shape is round, but in some of the longer bodied individuals it approaches the 
triangular form of most specimens of T. granarius. T. longicaudatus is similar in 
this character to T. granarius, but in T. australiensis a peculiar wide shape, with a 
slight emargination of the posterior margin, occurs in a few specimens. 

I have seen no specimens, and can find no records, of Lepidurus with triangular 
dorsal organs ; here the round shape seems to be usual but in most species it varies 
from round to oval (Text-fig. 4). In L. arcticus a peculiarly long, narrow oval shape 
is common and this does not appear to occur in other species (Text-fig. 4G). 

The growth rate of the dorsal organ is very strongly negatively allometric and 
this further complicates its use as a systematic character. In the larvae it is a 
relatively enormous structure so that in the first instar its median length is com- 
monly about half that of the carapace rudiment (Text-fig. 13B) . 

During growth to adult size its linear increase is only x 2 or x 3 while the relative 
growth of the carapace is naturally very much greater. This negative allometry 
appears to continue throughout growth. 

Linder made use of the relative arrangement of the eyes and the dorsal organ in 
his species of Lepidurus. He found that L. lynchi has the dorsal organ placed well 
behind the posterior boundary of the eyes and of the tubercles over the eyes, in 
contradistinction to the rest of the genus in which he found that the dorsal organ 
was placed in part between the eyes. L. batesoni sp. n. has the first arrangement 
(Text-fig 4D), but L. bilobatus — the other member of the long bodied group — has not. 

In Triops, almost all the specimens examined had the anterior margin of the 
dorsal organ between the eyes, the only exceptions being 8 of the 12 samples 
of T. australiensis examined, in which the arrangement was precisely similar 
to that of Lepidurus lynchi and L. batesoni (Text-fig. 4B) ; in the other four 
samples it was normal. This may indicate that not too much reliance should be 
placed on this character in either genus as a primary distinction between species. 

(8) Telson 

The telson bears an armature of spines on both dorsal and ventral surfaces, and 
variation of these have commonly been used in the past for separating species 
(Packard, 1883 ; Ghigi, 192 1; et at.). 

The important spines on the dorsal surface of the telson fall naturally into four 
groups to which it is convenient to apply names (Text-fig. 5) ; around the bases of 
the furca are rings of f ureal spines ; on the posterior margin of the telson of the 
larvae the first spines to appear are large and are identifiable in the adult — the 
posterior marginal spines ; the median area of the telson may bear a row of large 
spines or scattered smaller ones — the median spines ; around the dorsal sensory 
setae are rings or arcs of setal spines. 

The development, but not the origin, of these spines differs radically between the 
several species of Triops (Text-fig. 5), and the final arrangement is of the greatest 
importance in the systematics. 

All the specimens of Triops which were available were examined on a geographical 



basis, and no account was taken of previous determinations of the specimens ; it 
was found that there was a strong correlation between the spine pattern of the telson 
and the geographical distribution of the animals, but none with the sex or the 
number of segments. 

Fig. 5. Development of telson armature in Triops. a, larval stage (instar 2) common to 
all ; b, c, d, instar 5-6 ; b v c v t> v 3-4 mm. carapace length ; b 2 , c 2 d 2 , adults. B n , T. 
cancriformis ; c n , T. granarius ; D n , T. longicaudatus (the small arrows indicate the 

position of the posterior marginals in each case ; / 

spines) . 

furcal spines, m = median 

Without exception the specimens from Europe and western Russia have a small 
number of median spines arranged in an accurate row in the centre of the telson, the 
furcal spines are few and large, and the posterior marginals small, thin and remaining 
on the margin in the adult (Text-fig. 5A) ; this pattern occurs also in North Africa, 
the Middle East and northern India. In Africa south of the Palaearctic Region all 


the specimens have a larger number of small, relatively scattered medians in the 
mid-dorsal region, small, numerous furcal spines, and small posterior marginals 
(Text-figs. 5c, 6a). This pattern overlaps the European type in North Africa and 
the Middle East without forming intermediates, and then spreads across Central 
Asia to the Chinese coast. In North and South America the medians are similar to 
those of the European form, but there are two large spines, one on either side, at 
the posterior end of the median row, which represent the enormously enlarged and 
forwardly migrated posterior marginals, a fact which was confirmed by a study of 
the larval development (Text-fig. 5D). This form also occurs to the exclusion of 
others in the West Indies, the Galapagos, Oahu, and Japan ; a derivative occurs in 
New Caledonia (Text-fig. 16). 

Fig. 6. Telsons of Triops. a, T. granarius (trachyaspis-iorm) ; a v T. granarius ; b, 
T. australiensis ; c, T. cancriformis mauretanicus. 

The Australian forms have a very few small scattered medians, as in the African 
form, but they are so few in number as to be frequently absent (Text-fig. 6b) ; the 
rest of the armature is as in the African specimens. 

This gives a picture of four large groups which are to a great extent allopatric, 
but forming where they overlap no intermediates. The descriptions in the literature 
completely confirm this grouping and are too numerous to list with profit. 

As a systematic character this pattern appears to be perfect, there are no known 
intermediates, it is unlikely that it has any direct adaptive significance, and it is 
very easily seen in the specimens. 

It must be remembered, however, that although the pattern is stable the numbers 
of spines which form it are very variable ; in the pure line Triops cancriformis there 


were 1-4 medians, 2-4 furcals, and, once, small additional posterior marginals 
barely distinguishable from the primary pair. 

The number of these spines is fixed from a size of about 5 mm. carapace length, 
but as Text-fig. 5 shows, their relative sizes forming the true adult patterns are not 
stable until a little later, and allowance must be made for this (e.g. Apus mavliensis 

Fig. 7. Supra-anal plates of Lepidurus. a, L. apus apus ; a v L. apus lubbocki ; b, L. 
batesoni ; c, <j>, c v $, Lepidurus arcticus. 


Tiwari appears to have a different telson pattern from the other Indian forms, but 
this species was described on immature individuals in which the posterior marginals 
were still relatively very large). 

The spines encircling the dorsal setae are very similar in arrangement, though not 
in number, in all specimens of Triops examined, and no importance is attached to 

In Lepidurus, the spines corresponding to these four groups can be recognized, 
but they are to some extent modified ; the posterior marginals are the first to appear 
in the larval telson (L. apus) and the posterior margin of the telson on which they 
are borne expands rapidly to form the supra-anal plate, carrying them with it, and 
they are soon indistinguishable from the other spines on the margin of the plate in 
most specimens. The f ureal spines are similar to those of Triops granarius and do 
not differ much from species to species ; the setal spines are similar in form and 
lack of variation to those of Triops ; the medians form an elongate row along the 
mid-dorsal line of the plate. 

This median series shows considerable variation and has frequently been used in 
the separation of species ; the number of spines increases to a certain extent as the 
animal grows (Linder, 1952) ; and this process probably continues until the animal 
is at least 15-20 mm. in carapace length, to judge from a sample from Berlin of 
Lepidurus apus (BMNH) consisting of large and small individuals. 

The median spines may be borne on a slight keel, which is better marked in those 
specimens with a large number of spines in this series ; it is impossible to draw a 
line between presence and absence of a keel and contrary to Linder's opinion, I can 
make no use of it. 

The number of spines in the median series is useful in the systematics of Lepidurus 
species, serving to distinguish the nominate race of L. apus from the other three 
sub-species. L. arcticus has a much lower number than the rest of the genus. These 
differences are not connected with the relative size in the adults of the supra-anal 
plate ; both L. arcticus and L. lynchi have relatively low numbers of medians, 
although the former has the smallest supra-anal plate in the genus and the latter 
one of the largest. 

The marginal spines of the supra-anal plate vary in size and number and there is 
a connection between them and the median spines, when the latter are small and 
numerous, so are the former ; the marginals therefore are relatively large and sparse 
in Lepidurus arcticus, L. lynchi, and L. batesoni compared with the other species. 

The size of the supra-anal plate itself varies between species — the most obvious 
difference being in Lepidurus arcticus where it is very small — but a great deal of 
the observed differences are due to age and sex ; males (Text-fig. 7c) have relatively 
longer and more spatulate supra-anal plates than females (L. apus, Braem 1893 ; 
L. arcticus, Somme 1934) ; and the structure has a positively allometric growth rate 
throughout the period of growth (Braem, 1893 ; Campan, 1929). 

The end of the plate is occasionally incised medianly, giving it a bilobed appearance ; 
this appears in several species and has no importance (Linder, 1952), and it is to be 
expected that this will occur in species other than those in which it has already been 



The shape of the telson was used by Packard (1883), but Linder has shown 
that this is an unreliable character in Triops, though he records relative differences 
in length and breadth in some Lepidurus species. The variation is such that no 
reliance is placed on it here. 

The dorsal sensory setae are present, and similar in form, in all species of Notostraca 
that I have examined, and are longer in small than in large specimens. 

(9) Segmental armature 

Each segment which is exposed behind the carapace bears a series of spines on 
its posterior border ; this series is interrupted by the appendages and is continued 
vent rally only on the apodous segments. 

The form of these spines on the ventral surface of the apodous segments has 
frequently been used in the systematics of both genera ; Ghigi (1921) considered 
them to be important in Triops, as did Linder (1952) in Lepidurus. 


Fig. 8. Armature of apodous segments (a, b) and furca (c, d) of Triops. a, T. cancri- 
formis without supernumeraries ; b, T. granarius with supernumeraries (s) ; c, $ T. 
cancriformis or granarius ; d, <$, T. granarius. 

On the ventral surface of these segments the spines near the midline are usually 
smaller and set more closely together than the lateral ones. Linder (1952) considers 
that the number and relative size of these centrals is diagnostic of certain species ; 
he gives a variation of 13-28 teeth in Lepidurus packardi, L. couessi, and L. bilobatus, 
and only 8 in L. lynchi ; I find the variation so great that this character is effectively 
useless even though the differences are at times striking. 

In Triops, the shape of the marginal spines was used by Ghigi (1921), who 
found that they were squat in his T. mauretanicus, and finer in his other species ; 
I find that the European and North African material — which Ghigi divided into 
species — shows continuous variation in this character although, as he described, there 
is a tendency for the Moroccan specimens to have squat spines. 

These spines become rounded and heavy in males of Triops granarius in which 
the ventral armature of furca and telson are heavy ; they are thus heavier in males 
than in females of the same species. 

A more useful character on the apodous segments is the presence or absence of 
scattered supernumerary spines which occur between, and anterior to, the marginal 
spines and are much smaller than them (Text-fig. 8). In Triops cancriformis only one 



specimen has been seen which possesses any of these spines, a female from Palestine 
(coll. Goldschmidt) which had 3-4 small supernumeraries on one of the apodous 
segments. In the other species of Triops it is most unusual to find a specimen which 
does not possess at least a few supernumeraries on each apodous segment, and in 
the main there are 10-12 per segment. 
I have not seen these spines in Lepidurus. 

(10) Appendages 

The appendages have been described in considerable detail for one species (Triops 
cancriformis, Lankester 1881), but with the exception of one character have been 
little used in systematics. The endites of the first thoracic appendage are drawn 


o 1*0 

0*5 - 

5 IO 15 




Fig. 9. Relative growth rates in Triops cancriformis from Hampshire. Furca and 
longest endite of first thoracic appendage expressed as ratios furca/carapace and 
endite /carapace for various carapace lengths. Any slope on the line thus indicates 
allometric growth, since the growth rate of the carapace is isometric. 

out into filaments of which the longest endite (the fifth) forms an antenna-like 
structure in most forms. The length of the endite 5 and the number of segments of 
which it consists have been described by too many past authors to list. Even 
recently small differences in endite length have been considered to have significance 
(Tiwari, 1952). 
A study of the developing endite of Triops cancriformis and T. granarins has shown 


that it grows positively allometrically and that this rate continues, to some extent, 
throughout life (Text-fig. 9). So small differences in relative length are of no use 
in systematics without much more investigation of the growth rates of this structure 
in many populations. 

It is likely that the length of the endites is not related to the size of the animal 
per se, but to the instar that the animal has reached. Specimens which have grown 
rapidly, with a large increment at each instar would have a different endite length 
at a particular size from individuals with low instar increments due to poor condi- 
tions ; this is borne out by the great size and relative proportional differences 
seen in laboratory cultures. 

In museum samples, perhaps for this reason, the individual differences in this 
character appear to be greater than those to be expected between large and small 
specimens ; if a sample is arranged in order of carapace-length the endite /carapace 
ratio does not increase evenly up the series, as would be expected from the results 
obtained from the growth of single specimens, and the inference to be drawn must 
be that the specimens have grown at different rates and are of different sizes at 
each instar. 

This is further supported by the occurrence of two samples taken from the same 
locality in different seasons ; two such samples from a locality in the Saone area of 
France (MNHNP) collected two years apart consisted one of large, the other of 
small adult individuals, but the endite /carapace ratio was very similar in each ; in 
the first, individuals of 29-31 carapace length had a carapace /endite ratio of o*8o- 
0*84 ; in the second sample the same ratio occurred in specimens of 9-5-10*8 mm. 
in length. 

The situation is further complicated by the fact that there is sexual dimorphism 
in the endite length ; in 26 bisexual samples of Triops, 24 showed higher endite/ 
carapace ratio in the males. This may be due to the relatively shorter carapaces of 
males than to any real difference in endite length relative to the total size of the 
animals. In living specimens this appeared to be the case. 

It is very noticeable that the endites are shorter relative to the body length in 
both sexes of the longer bodied forms, but it is not possible to give reliable figures 
for this because of the parallel differences in the relative carapace length which 
renders inter-sample comparisons very difficult. 

The range of the endite /carapace ratio is very great within a species determined 
as such on other characters ; in Triops cancriformis this varies from 0*55-1*28, and 
in T. granarius from 0*47-1*52. No correlation with the geographical distribution 
could be found. 

In Lepidurus the endites are not as long as in Triops. Lepidurus arcticus and L. 
apus form a series from animals with very short endites which scarcely project 
beyond the carapace, to those in which they are relatively long. In L. batesoni the 
arrangement is unique ; the 5th endite of the first appendage is little longer than 
that of the second appendage, and the 6th endite forms a claw in both appendages 
although in all other Notostraca it is reduced to a small, soft lobe at the base of the 
5th endite in the first thoracic appendage, undergoing negative allometry during 
growth (Text-fig. 10). 



Linder (1952) remarks "... the legs of various species are known to be very 
similar to each other . . . ' ' ; I can find no reference to a comparative study of the 
appendages of Notostraca, and so it seemed valuable to attempt at least a preliminary 
survey. Unfortunately, it is necessary to dissect the appendages from the specimen 

Fig. 10. Thoracic appendages : a, first thoracic appendage of Lepidurus arcticus ; b, 
tip of second thoracic appendage of Triops cancriformis ; c, tip of first thoracic appen- 
dage of T. cancriformis ; d, first thoracic appendage of Lepidurus batesoni ; e, endite 6 
of first thoracic appendage of larval Triops cancriformis. (numbers = endites, 
fl = flabellum, br = bract.) 

in order to examine them properly, and this has naturally been possible with only 
a few specimens. Appendages of representatives of both sexes of all four species of 
Triops, and of several of Lepidurus were examined after mounting in polyvinyl 



The first antenna is present in all larvae and adults and is remarkably uniform 
in structure ; it bears at its tip three setae in all species that I have examined, 
although this number may be apparently altered by breakage. An aberration was 
seen in one specimen out of a sample of a trachyaspis form of Triops granarius from 
South Africa in which a pair of additional strong spines were present on one 

The second antenna must be present in all larvae, for in them it is the main 
locomotor organ ; the form and numbers of setae are identical in the larvae of all 
four species of Triops, but I have been unable to make comparisons in Lepidurus ; 
this appendage subsequently dwindles, due to negative allometry, and its locomotor 
functions are taken over by the thoracic appendages. It is often absent in large 


— ED 

Fig. 11. Mouth parts of Triops cancriformis to show second maxilla, (md — mandible, 
Mxi = first maxilla, Mxz = second maxilla, p = paragnath, T x = first thoracic 
appendage, ED = efferent duct of shell-gland). 

specimens, or it may be present (but very hard to detect) in the cleft in front of 
the mandible. I have found both conditions in large individuals of all species and 
can attach no importance to its presence or absence, contrary to the opinions of 
Linder (1952) and Spencer and Hall (1896). 

Ghigi (192 1 ) used the arrangement and relative sizes of the teeth on the triturating 
surface of the mandible as a systematic character in his division of European and 
North African material ; I have examined this character in many specimens con- 
specific with his material and there is so much variation that the invalidity of the 
character seems certain. Gauthier (1934) and Peres (1939) found similar variation 
in their North African material. 

The first maxilla has never been described comparatively ; I find the main varia- 
tion to be in the row of spines on the ventral edge of the appendage. These tend to 
be fewer in number and finer in Triops cancriformis than in the other species in the 


genus, but there is probably a complete gradation of intermediates, and I can make 
no use of the appendage in the systematics. 

The second maxilla has received more attention than the previous appendage and 
seems to provide an important character (Text-fig. n). Sars (1901) noted that 
the second maxilla was larger in Lepidurus than in Triops ; this is so, and this 
appendage is also larger in T. cancriformis than in the other species of the genus, 
but even in this species is so small that it can scarcely be of importance in the feeding 
mechanism. Linder (1952) reported that it was absent in large individuals of most 
species of Triops {australiensis, granarius, numidicus, longicaudatus) , but that no 
dwindling occurred during growth in T. cancriformis. I cannot agree with this. The 
second maxilla is present in all samples that I have seen of T. cancriformis and T. 
granarius, and absent in all those of T. longicaudatus and T. australiensis. Even in 
very small specimens of the latter two species it is absent ; in a narcotized female 
from Morawa in Australia it was clearly missing, yet this specimen was only 7-5 mm. 
in total length. 

In those species of Triops in which it is present it does not dwindle during growth ; 
in a T. cancriformis of 8*o mm. carapace length the second maxilla was 0*2 mm. long 
and in one of 18 mm. carapace length it was 0*4 mm. long ; in a T. granarius of 
7*5 mm. carapace length it was 0*13 mm. long, and in one of 13*8 mm. it was 0*33 mm. 

Thus, the presence or absence of a second maxilla provides a clear character 
separating two species groups within Triops, but it is present and well developed in 
all species of Lepidurus. 

The efferent duct of the maxillary gland arises at the base of this appendage, or 
from the position of the base if the appendage is absent ; this duct is longer in 
Triops than in Lepidurus (Sars, 190 1) and longer in male than in female Triops, in 
all the bisexual samples that I have examined. 

The endites of the first thoracic appendage have already been dealt with, and I 
could find no significant variation in the other lobes of this appendage, except 
that the flabellum was relatively larger in Lepidurus batesoni than in other species — 
another character of the 2nd thoracic appendage appearing on the 1st in this species. 

In the second thoracic appendage, the form of the terminal " claw " (= exopodite, 
Lankester 1881 ; apical lobe, Borradaile 1926 ; endopodite, Linder 1952 ; which 
I shall call endite 6 from the precisely similar ontogeny in the early stages of this 
and the other endites) was used by Ghigi (192 1) to differentiate between species of 
Triops ; I find great variety in the form of this lobe, but none that can be correlated 
with other characters. The relative lengths of endites 5 and 6 of this appendage 
show a sexual dimorphism ; in males endite 5 is commonly longer than endite 6, 
while in the females they are more nearly equal. This is well known and I have been 
able to confirm it in all the bisexual samples that I have seen. 

In the remainder of the thoracic series there is considerable variation in the shapes 
of the lobes of the appendages, but I can find none that are of importance between 
species ; however, all the specimens that I have seen of Triops granarius from 
Central and Eastern Asia have endite 6 of the mid-thoracic appendages of a more 
rounded shape than is usual. 

The armature of these appendages shows much variation in numbers of spines 


but not in their arrangement — a situation common in the armature on other 
structures in these animals. The numbers of spines in any group tend to increase 
during growth, but I have not been able to follow this closely. 

The abdominal appendages showed no differences which might be of use systemati- 
cally except to demonstrate once again a similarity between Triops cancriformis and 
the Lepidurus species ; the flabellum of Lepidurus bears a number (20-30) of setae 
around its outer margin, and these are reduced in number in Triops. But T. cancri- 
formis has a higher number (10-20) than the other species (e.g. T. granarius, 4-9). 

Linder (1952) mentions an inflated condition of the flabella in some specimens ; 
this I find to be due to post-mortem changes, especially in those animals which have 
died just after moulting, when the flabella are commonly so turgid with fluid after a 
few hours that they have the appearance of small red balloons. 

(n) Furca 

The length of the furca has been considered an important systematic character 
and was often included in even the early descriptions, but recent authors have thought 
it unreliable (Linder, 1952). 

The furca grow very rapidly in the larval stages of Triops but their positive 
allometry soon becomes less marked, though some relative increase probably occurs 
throughout life (Text-fig. 9). The furca are generally relatively longer in those 
specimens in which the endites of the first thoracic appendage are long, and are 
probably correlated in their development with the number of segments in the same 
way as the endites. That there is no direct correlation with the endite length is 
shown by Lepidurus arcticus in which the endites are very short, but the furca are 
similar in length to those of the other species. 

Gurney (1924) showed that sexual dimorphism in the armature of the furca 
occurred in Triops ; the spines on the ventral surface of the proximal region of the 
furca tend to be broader in males than in females, in extreme cases forming pro- 
tuberant scales (Text-fig. 8c, d). With few exceptions this dimorphism is more 
marked in long bodied forms, where the base of the furca tends to be relatively thick 
and rapidly tapering. In Lepidurus these spines are a little, but not much, thicker 
in males than in females. 


Oxyhaemoglobin has a characteristic absorption spectrum, and small differences 
in the wave-length at which the axes of the absorption bands occur have been 
demonstrated for several species of Daphnia by Fox (1945, 1946) ; similar 
differences in the position of the absorption band axes of the chlorocruorin of species 
and varieties of Sabella have been used systematically by the same author (1946). 

The blood of Notostraca contains a considerable concentration of haemoglobin 
in solution and the size of the animals is such that a volume of blood sufficient for 
spectroscopic analysis can readily be withdrawn by a micro-pipette. It seemed 
profitable to determine the wave-lengths of the absorption bands of as many 
populations of Notostraca as were available, and to apply the results to the 



A Hartridge reversion spectroscope was used to determine these wavelengths, in 
the manner described by Fox (1945), with a small refinement in technique to elimin- 
ate personal bias and errors due to parallax ; the operator kept his eye to the instru- 
ment throughout a series of measurements and an assistant made the readings on 
the micrometer head and noted them down. 

The wave-length of the axis of the oxyhaemoglobin-a band was determined by 
comparison with a sample of blood of similar optical density and of known wave- 
length (human blood, 5775 A). A number of readings were made on each sample of 
blood and the results were treated statistically. 

Several cultures of Triops cancriformis from different European localities were 
compared, using several adults from each culture. The results (Table I) show that 
although there is little individual variation, the three populations are apparently 
different from each other; these differences gave a probability of significance of 
0*02 when a t test was applied to them, and it may be taken that they are real and 
not referable to errors in the methods of measuring. 

Table I. — Wave-length of the axis of the oxy haemoglobin- cl band. Means of the readings 
taken on each animal, the averages of these means (Xj, the number of readings 
per sample(N), and the standard errors of the means ( ' S.E.J. The figures in 
parentheses indicate the distribution of the readings within a sample.* 

T. granarius 

T. longi- 


</ m \'t*tP /•/i*i/ , -vi fnv* 


m ij n n til c 

J. fv\jyo lslA/rbl/r bj \jr rri>vo 



6 il LtLLLll fA>5 










Means per ani- 

5779-7 (30) 

5778-3 (40) 

5777 -o (20) 

5776-7 (10) 

5778-2 (30) 

5777-3 (30) 

mal (A) 

5778-9 (30) 


5778-2 (30) 

5776-4 (30) 

5775-5 (30) 

5776-3 (30) 

5779 -o (30) 

5778-6 (20) 

5777-9 (30) 

5776-8 (30) 

5776-5 (30) 


5780-6 (30) 


5777-3 (30) 

5777*3 (30) 

5778-3 (30) 


5779-8 (28) 

5778-9 (39) 


5776-9 (30) 


x(A) . 





















This indicated that inter-specific differences must be considerably greater than 
these infra-specific ones to be of any value in systematics, and that many populations 
of a species must be tested before a value characteristic of that species could be deter- 

In fact, subsequent inter-specific comparisons gave results (Table I) very little 
different from those obtained with Triops cancriformis. 

The span between the axes of the oxyhaemoglobin and the carboxyhaemo- 
globin-a bands was also examined, for this is known to show differences between 
species in some cases (Fox, 1946). The results obtained with three species of Triops 
showed no differences which would justify further investigation, (span : T. cancri- 
formis = 50*1 A, T. longicaudatus = 48*2 A, T. granarius = 47*5 A). 

The results of both these investigations are of no practical value in the systematics 
of these animals — the differences being much smaller than those found and used in 
Sabella by Fox — but do serve to demonstrate that in this character, as in others, the 
species of Triops form a remarkably closely related group. 

* A single specimen of T. australiensis from Kalgoorlie was tested and gave a result of 5779*3 A. 




It has long been known that some populations of Notostraca contain both males 
and females, and some only females. The occurrence of large populations in which 
no males could be found has generally been assumed to indicate parthenogenesis, 
especially since it was also known that isolated females from such populations could 
produce viable eggs. 

Bernard (1889) found scattered testis lobes in the gonads of female Triops cancri- 
formis, Lepidurus apus and L. arcticus, and reported these occurrences as cases of 
hermaphroditism. Zograf (1906) found ovarian tetrads developing in the testis 

Fig. 12 Testis lobe in ovotestis of Triops cancriformis . 5(x section, Carnoy/Feulgen. 
(5 = sperms ; Sc = spermatocytes ; F.D.C. = follicle duct cells). 

walls of male L. apus, which must surely be an aberration of no use to the animal, 
for the oocytes subsequently degenerated. Spencer (1896) could find no testis lobes 
in the ovaries of Triops australiensis. 

Bernards' findings, largely ignored since his account, have been confirmed in the 
present work during investigation of the cytology of Triops and Lepidurus, and have 
been reported elsewhere (Longhurst, 1954). Hermaphroditism has been found in 
three species — Triops cancriformis, T. longicaudatus and Lepidurus arcticus — and it 
was found that females of the first two laid viable eggs in isolation and contained 
ovotestes (Text-fig. 12) ; no L. arcticus reached adult size. Females of Triops 
granarius and T. australiensis were unable to lay eggs in the absence of males, but 



did so readily as soon as males were put into the tanks with them and pairing had 
occurred ; this was presumably correlated with the fact that no testis lobes were 
present in the ovaries of the females of these species. 

There is no evidence to show that these latter two species of Triops are ever any- 
thing but bisexual ; in all the material I have examined of these species the large 
samples invariably contained both sexes, and there are no records in the literature 
of females occurring in large numbers without males. In T. cancriformis there appears 
to be a complex situation : in the southern parts of its range bisexuality is normal, 
and a female from Algeria had no testis lobes ; but in the more northerly regions 
males occur sporadically, often in very low proportions, so that some of the females 

Fig. 13. Notostracan larvae, a = carina of instar 3 Lepidurus apus ; b, metanauplius 
of Triops cancriformis ; c, Neonatus of Lepidurus arcticus. {do = dorsal organ.) 

probably reproduce without fertilization. In the extreme north of the range 
(Britain, Sweden) males are unknown and here females contain ovotestes. Unfor- 
tunately I have not been able to examine the gonads of females from a population 
with sporadic male occurrence. Mathias (1937) gives a review of the occurrences 
of males of this species. 

In Triops longicauddtus the geographical distribution of males is not so clear; 
however, specimens from Californian rice fields where males are unknown contained 
ovotestes, and Linder (1952) reports examining many specimens from the Galapagos 
Islands without finding males. Ueno (1935) records this species in Japan and found 
no males in 78 specimens. It may be that all the populations of this species from the 
Pacific region are hermaphrodite, for the small samples that I have seen from Hawaii 
(Oahu) and from New Caledonia contained no males. 



Reproduction in Lepidnrus is less well explored ; it is known (above) that herma- 
phrodite individuals occur in some species, but males of these are also known in some 
populations although their geographical distribution is not clear. The three long 
bodied species all appear to be bisexual. 

The size of the egg in Triops is variable, being larger in long bodied forms irres- 
pective of species ; with this is correlated a size difference in the mature follicles in 


6 9 



Fig. 14. Notostracan chromosomes, a-c, 2 spermatocyte telophase ; d-e, young 
oocyte ; f, resting unfertilized ovum ; g, spermatocyte diakinesis. a, Triops longi- 
caudatus ; b, T. australiensis ; c, d, e, T. granarius ; f, T. cancriformis ; g, Lepidurus 
apus. (g is half the scale of the remainder and is redrawn from Goldschmidt, 1953.) 

the ovary : T. cancriformis (33 segments), 0*28 mm. ; T. granarius (38 segments), 
0*45-0*52 mm. This size difference is correlated with size differences in the resultant 
eggs and larvae, the long bodied forms having the largest larvae. In Lepidurus the 
differences may be even more marked and are diagnostic of species in at least one 
case ; the larvae of L. arcticus hatch at a later stage of development than those of 
the other species (Text-fig. 13), and the eggs are very much larger, the ovarian 

ZOOL. 3, 1. 3 



follicles are relatively few in number, large and elongated in form, and completely 
fill the space above the gut. 


Moore (1893) was the first to examine the Notostraca cytologically, and from obser- 
vations on the divisions of somatic cells he concluded that Triops cancriformis was 
amitotic (zn = 1). 

While the present work was in progress, Goldschmidt (1953) recorded chromo- 
somes in the testes of Lepidurus sp. from Palestine, finding that aceto-orcein squashes 
gave diakinesis stages where n = 6 (Text-fig. 14G). I have since determined her 
material as L. apus lubbocki. 

Using testis smears I found that two populations of Triops granarius from South 
Africa showed the number n = 4 ; this was most clearly seen in the telophase stages 
of the spermatocyte (Text-fig. 14c). This number was confirmed in females of the 
same populations in which a haploid set of chromosomes was readily seen in the very 
early oocytes (Text-fig. 14D, e). 

Sections of the ovotestes of Triops cancriformis from the European cultures showed 
n = 4 in the oocytes, most clearly in the late oocytes or ova, resting in the longi- 
tudinal oviduct (Text-fig. 14E) ; squashes of the same gonads showed n = 4 in 
secondary spermatocyte telophases in the testis lobes. 

In both of the above species of Triops the diploid number, 2n = 8, was confirmed 
by finding mitoses in cells of the expanding follicle walls, at the stage of increase in 
size of the oocyte and nutritive cells. In T. longicaudatus from California the number 
n = 4 was seen in early oocytes (Text-fig. 14A) and in spermatocyte telophases, but 
it was not possible to confirm the diploid number. 

A single male of Triops australiensis from Kalgoorlie was available and testis 
smears were made from it which showed, in the spermatocyte telophase, a comple- 
ment of n = 5 ; this number was counted in at least 16 nuclei. The 5 chromosomes 

Table II. — Chromosome numbers of Notostraca 




Diploid number 




L. apus lubbocki . 




T. c. cancriformis 




(2 spermatocyte 

(resting oocyte) 

(follicle duct 

T. granarius 

T. 1. longicaudatus 

T. a. australiensis 


(2 spermatocyte 



(2 spermatocyte 



(2 spermatocyte 


(young oocyte) 

(young oocyte) 

(follicle duct 


in this haploid set are all sub-equal in length (Text-fig. 14B), and it seems probable 
that this situation could have arisen by the fragmentation of one chromosome in a 
set of four, such as the other species possess, in which one is nearly twice as long as 
the other three. 

My findings and those of Goldschmidt are summarized in Table II, and both agree 
in ascribing a low number of chromosomes to the Notostraca so far examined. 
Moore's account of amitosis may be true of the cells that he studied, but even 
that is doubtful in view of my chromosome counts in epithelial cells ; his inference 
of a single chromosome was obviously incorrect. 

All the Triops material was fixed in Carnoy's or Susa's fluids and was stained with 
the Feulgen reaction. 


Very many accounts of the ecology of the Notostraca and of their appearance in 
temporary pools have been published, and a clear picture of their biology can be 
derived from the literature. 

There appear to be almost no ecological differences between the species of Triops, 
a slight difference between Triops and most species of Lepidurus, and a marked 
divergence between L. arcticus and the rest of the Notostraca. 

So far as is known, Triops occurs only in waters which dry out regularly, and the 
eggs normally hatch in the field only after a period of dessication. The habitat 
itself may vary a great deal in size from tiny rain pools and cart-ruts to large tempor- 
ary lakes (Main, 1953) and the water may be fresh as in the Hampshire locality 
(Hobson and Omer-Cooper, 1935), brackish as in the Scottish pools (Balfour-Browne, 
1909), or saline as in the lakes on the Tibetan plateau (Schlagintweit, 1872) and in 
Australia (Main, 1953). 

The farming practice of rice fields makes these ideal situations for Triops, which 
are sometimes present in enormous numbers and have been recorded as a rice field 
pest from many parts of the world (they uproot the rice seedlings) . 

Lepidurus may occur in temporary pools, but there seems to be a tendency for 
most species to live in waters which dry out less regularly than the Triops pools. 
Lundblad (1920) records that Lepidurus apus is a spring form which appears when the 
temperature rises early in the year in ponds or ditches which have held water all the 
winter : Linder (1952), on the other hand, says that he has seen the same species 
in Sweden in temporary pools. L. apus viridis occurs in a pool near Christchurch, 
New Zealand, which dries out in normal years, but is only very rarely found in neigh- 
bouring permanent ponds (G. Parry, personal communication). Lepidurus spp. 
regularly occur in temporary, often alkaline, pools in North America (J. Lynch, in 
litt.) . L. Glauert has sent me a map showing all the records for Triops and Lepidurus 
in Western Australia and it is clear that Lepidurus is restricted to the south-western 
coastal belt where there is regular winter rain and Triops to the arid interior where 
rainfall is, at the most, sparse. 

Lepidurus arcticus is confined to the boreal and alpine regions of the Holarctic, 
where it may occur in large lakes and form an important food of Salmonidae and 
where the eggs could never be dessicated (Somme, 1934), it is also in pools which 


normally do dry out each year (J. Mohr, in litt.). The only other records of Lepi- 
durus occuring in lakes refer to L. lynchi (Linder, 1952). 

It has been thought that there is a difference between the two genera in the con- 
ditions necessary for the hatching of the eggs : Fritsch (1866) and Grasser (1933) 
thought that Triops eggs could hatch only after dessication, while Brauer (1877) 
believed that the eggs of Lepidurus were incapable of withstanding dessication. 
Schaeffer (1756) and Kozubowsky (1857) showed that the eggs of Triops were some- 
times capable of hatching without drying out. This has been confirmed in the present 
work and it is now known that the eggs of at least three species of Triops can hatch 
out, after an interval for development, in the water in which they were laid, or will 
remain viable if dried out and will hatch when replaced in water. This accounts 
for the report by Mathias (1937) that Triops has two types of eggs : the one drought 
resistant and the other not. 

Fox (1949) reported that the eggs of Lepidurus apus viridis were able to hatch 
after drying, and I have confirmed this with another mud sample from the same 
locality as his ; L. arcticus from Iceland were also hatched from dried eggs. The 
ecology of Lepidurus outlined above indicates that the eggs of this genus must in 
many cases be drought resistant. 

The eggs of Triops probably do not often hatch out without prior dessication, 
as the water must have a low osmotic pressure for this to occur, a condition unlikely 
to be found on the bottom of a pool where the eggs are laid, but most likely when the 
pool refills with water after rain and the eggs float to the surface. 


From the analysis of the characters used in the systematics of the Notostraca 
it is evident that many of the obvious differences between individuals must be 
correlated with differences in the number of body segments (Text-fig. 2) ; in Triops 
there is no discontinuity in the variation of this number, so that neither the number 
itself nor characters correlated with it can validly be used in the separation of species. 

In this genus, as the number of segments increases to give a long bodied form so 
the carapace becomes smaller, rounder and flatter ; the number of legs remains 
about the same and consequently the apodous number becomes higher ; the furca, 
and the endites of the first thoracic appendage become relatively shorter ; the dorsal 
organ becomes larger, less elevated, and more often triangular in outline ; the sulcus 
spines become smaller and more numerous, and the terminal spine of the carina is 
more frequently absent ; sexual dimorphism becomes more marked. 

Similarly, certain characters are correlated with the sex of the specimen ; males 
tend to have more segments than females and so the characters outlined in the previous 
paragraph vary from male to female within a population ; characters other than 
these also show sexual dimorphism : the efferent duct of the shell gland is longer 
in males than in females ; endites 5 and 6 of the second thoracic appendage are more 
nearly equal in length in females ; the nth thoracic appendage bears a brood pouch 
in females ; the ventral armature of the furca of males is coarse, often forming scales 
rather than spines, and with this is correlated the ventral armature of the telson 
and apodous segments, which vary in the same way between males and females. 


These two considerations then — segment number and sex — invalidate a number of 
characters in the systematics of Triops. With those that remain there appear to 
be good grounds for dividing the genus into four groups, each with a geographical 
basis. The valid characters appear to be : the armature of the telson, the presence or 
absence of the second maxilla, and the arrangement of the eyes and dorsal organ. 
Of these the most important is the armature of the telson, which is diagnostic for 
each group ; the other characters, together with a few less well-marked ones, confirm 
this primary grouping. 

Two groups are partially sympatric, and here there is no tendency to hybridize 
and no intermediates have been found in the areas of overlap ; there is a record, 
moreover, of two species — which correspond to two of these groups — living in the 
same pools in several localities in Morocco (Peres, 1939). These two forms, therefore, 
behave precisely as biological species (sensu Mayr, 1942) would be expected to do 
when they become sympatric ; it seems justifiable to consider them, therefore, as 
species and since the degree of morphological difference between all four groups is 
similar to that found between these two, then all four may equally well be regarded 
as species. 

Each species, then, has a geographical basis and a clear cut character in the telson 
armature, but contains populations which are remarkably different in general appear- 
ance depending on the number of body segments in the specimens. What determines 
the body length is completely obscure, for there appears to be a general tendency 
for the longer bodied animals to occur in the warmer regions of a species range, but 
there are many blatant exceptions to this generality ; Barnard's Apus ovamboensis 
is a population of exceptionally short bodied Triops granarius from a very hot and 
dry part of Africa. 

It is likely that an experimental study of the effect of environmental factors on 
morphology would throw some light on this problem, for perhaps the effects are seen 
only after a considerable number of generations. 

Several species show some indications of the presence of geographical races ; the 
differences between these are much slighter than the specific differences, and their 
distribution indicates that the geographical barriers between them are very slight. 
Triops cancriformis will serve as an example ; the populations in Morocco and 
southern Spain differ in several respects from the rest of the species, even though 
in Spain there is no geographical barrier where the change occurs, and there is some 
evidence that intermediates occur where the races meet, for in Seville and near Gib- 
ralter the specimens are identical with the Moroccan ones and from Valencia they are 
similar to the specimens from the rest of Europe, but in Ciudad Real — midway 
between the two areas — the specimens are intermediate. A similar zone of inter- 
mediates seems to occur in Spanish Morocco east of Ceuta. 

Ghigi (1921) considered these races of Triops cancriformis to be species and des- 
cribed them as such ; Colosi (1922) and Peres (1939) referred to Ghigi's species as 
varieties of the single species T. cancriformis but Gauthier (1934) used a true 
trinomial nomenclature and considered them to be sub-species or geographical races ; 
I follow this nomenclature and believe them to be sub-species in the sense that 
Huxley (1942) and Mayr (1942) used the term. 


The arrangement of the species seems to follow different principles in Lepidurus 
from that in Triops, and the following account owes very much to the work of Linder 
(1952) who demonstrated the existence of two species-groups within the genus, each 
group characterized by its segment numbers. 

The variation in segment number in Lepidurus is not so great as in Triops, and 
many characters show no correlation with this number in Lepidurus though the 
correlation is obvious in Triops : the armature of carina and sulcus ; the endites of 
the first thoracic appendage ; the number of apodous segments and the size and shape 
of the dorsal organ. The segment number shows a marked discontinuity at about 
30 ; the vast majority of specimens have less than this number, and those which 
have more are aberrant in other respects. 

Within the short bodied group there is relatively little differentiation, but Lepi- 
durus arcticus can be separated at once, and is distinguished by its range, its habitat, 
its relatively short supra-anal plate and endites. The remainder of the group appears 
to comprise a single species, with a few rather ill-marked, but very widespread, sub- 

Linder (1952) showed that Lepidurus couesii was conspecific with specimens deter- 
mined as L. macrourus Lilljeborg. I can find no differences between the specimens 
that I have seen of either of these species and those of the European L. apus. The 
ranges of variation in the segment number are similar, the size and armature of the 
supra-anal plate is the same, and so is the armature of the carapace. 

In California a form occurs which is admitted as a species by Linder, Lepidurus 
packardi ; in these specimens there is usually a slightly higher number of segments 
than in the normal L. couesii (= L. apus), the apodous number tends to be higher, 
and the sulcus spines are very small, numerous and closely packed. In the Medi- 
terranean region there occurs a form which diverges from typical L. apus in a similar 
way except that the sulcus spines are normal. Both of these forms replace the typical 
race over very restricted areas, but there appear to be no effective geographical 
barriers separating the several ranges from one another, and for these considerations 
I propose to consider these forms to be sub-species of the typical and widespread 
L. apus. 

The Australasian forms do not differ from typical Lepidurus apus as much as do 
the above two sub-species, and many specimens would be indistinguishable if placed 
in a sample from Europe. However, I have seen no specimens which have the high 
number of central spines on the supra-anal plate which are very common in the 
typical L. apus, so that there is a slight degree of morphological differentiation and 
I propose to consider the Australasian form as another sub-species, L. apus 
viridis. The South American forms which I know only from descriptions must 
be considered as another sub-species of L. apus. 

The rest of the genus comprises the few longer bodied specimens which are known. 
All have coarse and sparse marginal spines on the supra-anal plate — a character 
which they share with Lepidurus arcticus — and a low number of central spines on the 
same structure ; specimens from three localities bear uniquely large spines on the 
carina and the carapace margin and these are Linder's L. lynchi, typical form and 
var. echinatus. A small sample from Russia has a considerably higher apodous 



number than the rest of this group and shares with L. lynchi a peculiar arrangement 
of the eyes and the dorsal organ, which resembles the arrangement in Triops austra- 
liensis where the anterior margin of the dorsal organ may be placed well behind the 
posterior margin of the eyes. A single specimen from Utah was placed in this 
species-group by Linder on account of its segment number and seems to agree with 
the description of Lepidurus bilobatus Packard, having a normal carapace armature 
and normal arrangement of the eyes and dorsal organ. The only possible arrange- 
ment at the moment is to consider all these three forms as three separate species, 
but it seems very probable that future work will be able to find a connection at least 
between L. bilobatus and the Russian sample (L. batesoni sp. n.) and perhaps between 
these and L. apus. 

It will be noted from the foregoing account how widespread are the species of the 
Notostraca and this is probably accounted for, as it is in some other invertebrates, 
by their passive distribution ; the dried viable eggs must be blown around by wind, 
and transport by birds is not unthinkable, for the eggs when laid are extremely sticky 
and remain so for some days while the shell hardens, and so could presumably adhere 
to larger animals, The eggs of other phyllopods are known to be capable of passing 
unharmed through the guts of amphibia (Mathias, 1937), and birds are known to 
eat Notostraca ; starlings (Decksbach, 1924) and gulls (Balfour-Browne, 1909) 
are. recorded as feeding on Triops cancriformis, and Summerhayes and Elton (1923) 
watched Arctic terns feeding Lepidurus arcticus to their young, and thought that 
they might drop them accidentally into fresh pools on their way to the nest. 

A passive distribution such as this must mean that geographical barriers are not 
nearly so effective as they are for sedentary, or non-passively distributed animals, 
and has produced species with world-wide distribution in other animals, such as 
Tardigrades and Rotifers. 

In addition, the group has a very long geological record and has had ample time 
to occupy all suitable areas ; fossils from the Permian (Guthorl, 1934) are clearly 
Notostracan carapaces, and forms from the Triassic of Europe (Trusheim, 1938) 
are certainly Triops and differ from extant T. cancriformis only in the small size 
of the terminal carinal spine. The upper Triassic L. stormbergensis (Barnard, 1929) 
from South Africa is very similar to recent species except that the supra-anal plate 
has, apparently, no central spines as in some extant L. arcticus. 

It is impossible from these few fossils to give any account of the history of the 
group, except to point out how little evolution has occurred in the space of 170 
million years since the Triassic forms were alive. 

However, something can be deduced about more recent changes in distribution. 
During the Pleistocene glaciations Lepidurus arcticus was much more widespread 
in western Europe than it is now ; this species is known from lacustrine beds of 
that time from Scotland (Bennie, 1894) and from the Isle of Man (Geikie, 1894), and 
now shows in Scandinavia the typical distribution of a boreo-alpine relict occurring 
at sea level in the North and at progressively greater altitudes towards the South 
(Somnie, 1934). 

Triops cancriformis must have been absent from much of its present range at the 
same period, and the post-glacial extension may have been performed largely by 


hermaphrodites, which predominate in the northern parts of its range ; for these 
would be more efficient in dispersal as only a single egg would be required to effect 
a colonization. Perhaps the increasingly sporadic occurrence of males towards the 
north indicates a spread northwards of bisexuality. 

All the known rice field populations consist entirely of females (or hermaphrodites 
if parthenogenesis is assumed not to exist in these animals) and this strengthens 
the above argument, for these are relatively new habitats for Notostraca, and a coloni- 
zation by hermaphrodites ahead of the bisexuals seems to have occurred. 

The westward extension of Triops longicaudatus across the Pacific from the largest 
area of distribution in North America, may also be of relatively recent origin and 
has apparently been performed again by hermaphrodites, for no males are known 
from the Pacific populations (p. 47). 


Keilhack (1909) and Fox (1949) have shown that the generic name Apus Schaeffer, 
1756 should be rejected in favour of Triops Schrank, 1803 ; this practice will both 
accord strictly with the Rules of Nomenclature and will avoid further confusion in 
this genus, and in the avian genus Apus Scopoli, 1777. I therefore propose to follow 
Keilhack in the use of Triops. 

The use of trinomials is necessary to describe formally the geographical sub-species 

The lists of synonymies before each species are not complete lists of references to 
that species, for these would be too long and not very useful — but include (a) the 
names, and the various spellings of each name, that have been applied to that species, 
and (b) important descriptive works. 

Apus has been applied to Crustacea other than Notostraca ; Apus pisciformis 
Schaeffer is an Anostracan and Apus caudatus De Kay, was a crustacean parasitic 
on a crab. 


The following key should serve to identify specimens down to species, but the 
user must bear in mind the extent of variation in any character and not expect the 
figures to match the specimen in details of armature. It would be very valuable if 
the segment number, the apodous number, and the number of appendages were 
given in any future records. 

It must also be remembered that every individual in a sub-species may not be 
typical of it ; many ornithologists consider a sub-species to be valid if 75% of its 
specimens can be placed in it without question. In Triops cancriformis , for example, 
it may be difficult to ascribe single specimens to the nominate race or to T. cancri- 
formis simplex, for the smooth carina which is typical of the latter occurs in some 
individuals of the former, but I have no knowledge of whole samples of the nominate 
race with smooth carinas. 

Key to species of Notostraca 

1. Supra-anal plate present (Text-fig. 7) . . . . . . . (Lepidurus) 2. 

Supra-anal plate absent .......... (Triops) 6. 


2. Segments more than 30 .......... . 3. 

Segments less than 30 ........... 5. 

3. Anterior margin of dorsal organ between eye tubercles (Text-fig. 4E) L. bilobatus (p. 53). 
Anterior margin of dorsal organ well posterior to eye tubercles (Text-fig. 4D) . . 4. 

4. Carina and/or lateral margins of carapace with large spines. (Text-fig. 3) Apodous 

segments 3-5 . . . . . . . . . . L. lynchi (p. 53). 

Without large spines on carina or lateral margin. Endite 6 of first thoracic appendage 

as in Text-fig. iod. Apodous segments 8-9 .... L. batesoni (p. 54). 

5. Supra-anal plate short, 0-5 spines centrally, marginals few (Text-fig. 7c). Endites 

scarcely project beyond edge of the carapace . . . . . L. arcticus p. 52). 

Supra-anal plate long (Text-fig. 7 a, b), 4-100 central spines, marginals numerous. 
Longest endites with 3/4 of their length projecting beyond edge of carapace (Text- 
fig. 2i) . . . . . . . . . . . L. apus (p. 50). 

6. Second maxilla absent ........... 7. 

Second maxilla present (Text-fig. 11). . . . . . . . . 8. 

7. Posterior marginals sub-equal to medians and well forward of the margin (Text-fig. 

5D 2 ). Medians large, 1-4 in number, in a row T. longicaudatus (p. 46). 

Posterior marginals reduced and marginal. Medians small, scattered when more 

than 3-4, often absent. (Text-fig. 6b) . . . . . T. australiensis (p. 48). 

8. No supernumerary spines on the apodous segments (Text-fig. 8a) . Medians large, 1-4, 

in a row (Text-fig. 5B 2 ) . . . . . . . . T. cancriformis (p. 41.) 

Supernumerary spines present on apodous segments (Text-fig. 8b). Medians of 
various sizes, scattered except when less than about 5 are present, when they form 
an irregular row (Text-figs. 5C 2 , 6a, a x ) . . . . T. granarius (p. 44.) 

Genus TRIOPS Schrank, 1803 

1756. Apus Schaeffer (in part). 

1758. Monoculus Linn, (in part). 

1803. Triops Schrank. 

192 1. Thriops (sic.) Ghigi. 

1 92 1. Proterothriops Ghigi. 

Triops is at once separable from Lepidurus by its lack of a supra-anal plate ; even 
if a structure resembling a rudimentary plate (Text-fig. 7c) is found in some specimens 
of Triops cancriformis (Linder, 1952) there is never any doubt as to which genus a 
specimen belongs. 

Proterothriops was erected by Ghigi for the reception of the long bodied forms of 
the genus ; he was struck by the great difference in general appearance between 
long and short bodied forms which I have now been able to show to be conspecific 
in a number of cases. 

Binoculus Geoffroy, 1762 and Apodis Zaddach, 1841, are not in a strict binomial 
system and are ignored on this account. 

Triopes Schrank, 1803, appears later in the publication than Triops and must be 
regarded as a spelling lapse. 


1756. Apus cancriformis Schaeffer (in part), 
1758. Monoculus apus Linn, (in part). 
1801. Apus cancriformis Bosc. 



Apus viridis Bosc. 

Triops palustris Schrank. 

A pus montagui Leach. 

Apus himalayensis Packard. 

Apus halicienis Fiszera. 

Apus lublinensis Fiszera. 

Apus varsovianus Fiszera. 

Triops cancrifovmis (Bosc) Keilhack. 

Apus cancrifovmis bidens Sidorov. 

Apus carter if or mis transcaucasicus Sidorov. 

Thriops simplex Ghigi. 

Thriops mauretanicus Ghigi. 

Thriops apulius Ghigi. 

Thriops cancriformis var simplex, Colosi. 

Triops cancriformis simplex (Ghigi, 1921) Margalef. 

Triops cancriformis mauretanicus (Ghigi, 1921) Margalef. 

Type. Original is unknown, and designation of neotypes desirable. A sample 
from Kirkudbrightshire is selected (BMNH, 1907.10. 17. 1-4) and consists of a 
number of neoparatypes. 

Range. Western Europe (Spain to Sweden) east to Russia ; North Africa, 
Balkans, Asia Minor, Middle East to India. Individual records too numerous to 
list ; Lundblad (1920) gives many for Europe, Decksbach (1924) many for Asia. 
Does not extend beyond 6o° N and range in Russia obscure, but no records authenti- 
cated for Eastern Asia. 

Habitat. Temporary fresh or brackish waters ; occurrence depends on the pools 
filling when the temperature is high enough for development, so usually summer form 
in Europe, spring form in N. Africa. 

Rice field pest in N. Italy, Spain and La Carmargue. 

Description. Head. — Dorsal organ round, oval or rarely triangular, small and 
with its anterior margin between the eyes. Second maxilla present in all specimens 
examined and relatively larger than in the rest of the genus. 

Carapace. — Shape generally oval, more round in males. Carina with terminal 
equal to, or longer than, sulcus spines (Text-fig. 15) ; sometimes number of smaller 
spines on posterior carina. Sulcus spines long, 24-44, usually about 30. 

Body. — Segments 32-35 in both sexes ; apodous 4-7 in ?£, 5-9 in <$<$. Apodous 
segments without supernumerary spines on ventral surface. 

Telson. — Median spines large, 1-4, in a single median row (Text-fig. 5b). Furcal 
spines large, number sometimes varying from side to side of a single animal, and with 
few scattered spines on lateral face of telson anteriorly. 

Posterior marginals small, fine and marginal in the adult, and posterior margin 
of telson sometimes drawn out to resemble a rudimentary supra-anal plate (Text- 
fig. 6c). 

Furca generally long. 

Appendages. — 48-57 pairs recorded, but variation probably greater. Endites of 
first thoracic appendage long. 

Sexual dimorphism. — Not as well marked as in other species ; no males with furcal 
scales seen. Abonyi (1926) records two sets of males in one summer in the same pool 



of which the second batch had very reduced sexual dimorphism — so a dimorphism 
of males may exist ? 

Larvae. — Metanauplius (Text-fig. 13) ; dorsal organ of instar 1 generally round. 

Reproduction. — Bisexual and hermaphrodite ; in the northern regions no males 
occur and such females as have been examined are hermaphrodite. In the South 
populations are bisexual and in central Europe males occur sporadically. Mathias 
(1937) reviews occurrence of males. 

Geographical Races 
1. Triops cancriformis cancriformis (Bosc) 

Range. Whole of species range with exception of that occupied by the other 
two sub-species. 

Description. Carina bears 0-10, generally 2-3, small teeth in front of the 
terminal spine (Text-fig. 15E) and no large samples without specimens showing 
these spines are known. Furcal spines small (Text-fig. 5B 2 ). Dorsal organ round- 

Hermaphrodite and bisexual. Chromosome number zn = 8. 


Fig. 15. Carinas of Triops. a-c, T. granarius ; d, T. cancriformis mauretanicus ; e, 
Ej, T. c. cancriformis ; f, T. cancriformis simplex. 

2. Triops cancriformis simplex Ghigi 

Range. North Africa, Ceuta to Egypt. 

Description. Characters of Thriops simplex Ghigi. Carina quite smooth in 
front of the terminal spine (Text-fig. 15F) ; this is invariable in the specimens I 
have seen from this area, and in those seen by Gauthier (1934), Colosi (1922) and 
Ghigi (1921). Furcal spines small (Text-fig. 5B 2 ). These populations frequently 
show a tendency to a general weakness in the strength of the armature and include 
the sample from Kebili (p. 14) which has specimens with no carapace armature 
whatever. The terminal spine of the carina is frequently reduced. 


Apodous segments frequently higher in number than the nominate race ($? 5-7, 
instead of 4-6). 

3. Triops cancriformis mauretanicus, Ghigi 

Range. N.W. Africa : French Morocco and Tangier. S. Spain and Balearics 

Description. With the characters of Thriops mauretanicus Ghigi : armature 
very strongly developed ; carina with a number of teeth posteriorly (Text-fig. 15D), 
the largest often sub-equal to the terminal spine. Furcal spines very large (Text- 
fig. 6c) ; ventral marginal spines on apodous segments very coarse. Apodous 
number is similar to that of sub-species 2. Dorsal organ oval. 



Apus viridis Bosc was applied to figures of Schaeffer (1756) which showed juvenile 
Triops cancriformis ; Triops palustris Schrank is presumably of this species because 
of its N. European locality ; Apus montagui Leach, type in BMNH and clearly of 
this species ; Apus himalayensis Packard, author's figures show clearly the cancri- 
formis-type telson ; Apus varsovianus, etc. Fiszera, figures similarly show cancri- 
formis-type telson ; Sidorov's two sub-species are insufficiently described and can 
both be referred to this species ; Thriops apulius Ghigi ; this could be either sub- 
species 1 or 2 as it was a single specimen with a smooth carina, but from its Italian 
locality is probably of sub-species 1. 


1864. Apus granarius Lucas. 

1865. Apus numidicus Grube. 
1877. ? Apus dispar Brauer. 
1877. Apus sudanicus Brauer. 
1880. Apus dukeanus Day. 
1886. Apus bottegoi Prato. 

1886. Apus namaquensis Richters. 

1893. Apus sudanicus var. chinensis Braem. 

1893. ? Apus sudanicus var. braueri Braem. 

1893. Apus numidicus var. strauchii Braem. 

1893. Apus numidicus var. dybowskii, Braem. 

1895. Apus somalicus Wedenissow. 

1899. Apus bottegoi, Bouvier. 

1899. Apus trachyaspis Sars. 

1899. Apus sculleyi Sars. 

1899. Apus namaquensis Sars. 

1899. Apus granarius Sars. 

1907. Apus elongatus (nom. nov. for namaquensis Sars) Thiele. 

1920. Apus zanoni Colosi. 

1921. Apus asiaticus (nom. nov. for granarius Sars) Gurney. 


1922. Triops uebensis (nom. nov. for bottegoi Bouvier) Co'osi . 

1924. Apus ovamboensis Barnard. 

1927. Apus numidicus var. sinensis Ueno. 

1929. Apus cancriformis , Barnard, (non Schaeffer, 1756). 

1929. Apus sudanicus, Barnard. (? Brauer, 1877.) 

1934a. Apus granarius, Gauthier. 

1937. Apus sudanicus, Gauthier. (non Brauer 1877). 

1939- Apus numidicus, Peres. 

1940. Apus sinensis Ueno. 

1952. Apus mavliensis Tiwari. 

1952. Apus orientalis Tiwari. 

Type. MNHNP ; $ Holotype, unregistered other than label " Type A. granarius 
Lucas ". 

Range. South Africa to China ; the whole of Africa except for unsuitable forest 
regions in west and centre ; Middle East, India, central and eastern Asia to the north 
Chinese coast. Single sample in MNHNP labelled " near Paris " almost certainly 
mistake in labelling. 

Habitat. Temporary fresh and brackish waters ; unknown from rice fields ; 
Peres (1939) compares its distribution in North Africa with that of Triops cancri- 
formis and finds latter in steppe zones (300 mm. annual rain) and in the sub-steppe 
(300-500 mm.) while present species is restricted to the sub-steppe. 

Description. Shows great variation in body length and includes very short and 
long bodied forms with all the associated variation. 

Head. — Dorsal organ triangular, oval or round in the short bodied forms, anterior 
margin between the eyes. Second maxilla present in all specimens examined except 
in one case (see below). 

Carapace. — Shape variable, round to oval. Carina with terminal spine in short 
bodied forms only (Text-fig. 15), generally smooth. Sulcus round (or squared in 
some African short bodied forms), spines many, often reduced in size, 24-72. 

Body.— Segments : $$, 32-42 ; <?<?, 32-43- Apodous : ?? 4-13 ; £<£, 6-14. 
Apodous always with varying number of supernumerary spines vent rally (Text-fig. 

Telson. — (Text-fig. 5C 2 , 6a). Medians small, numerous, scattered, but when low 
in number may form a rough row medianly, though this never as accurate as in 
Triops cancriformis, and spines smaller. Furcals small and with many scattered 
in front of them. Posterior marginals small, squat and marginal. Furca variable 
with body length. 

Appendages. — 44-46 recorded, but variation probably much greater than this. 
Endites of first thoracic appendage variable with body length (Text-fig. 2). 

Sexual dimorphism. — Strong in many long bodied specimens, males having strong 
scales on furcal (Text-fig. 8d). 

Larvae. — Metanauplius, those seen having in instar 1 a trapezoid-shaped dorsal 
organ which later changes to triangular. 

Reproduction. — Bisexual. Chromosome number 2n = 8. 

46 a review of the notostraca 

Geographical Races 

With present knowledge not possible to recognize any with certainty. Some indi- 
cations known — many South African (not S.W.A.) and Middle East (Bombay- 
Baghdad) specimens have reduced sexual dimorphism even in long bodied forms ; 
these correspond to Apus asiaticus Gurney and this name may eventually be applied 
to a race with a restricted range. Some specimens from Eastern Asia have reduced 
second maxilla, and this is even absent in few out of large sample (BMNH 1935. 
6.18. 7-12) and these might be separated off as another race, but evidence too weak 
at present. 


Apus sudanicus Brauer belongs here doubtfully as the description is inadequate 
and I have not seen the types, but Barnard's specimens determined as this are cer- 
tainly Triops granarius. A number of species are placed here on evidence of original 
descriptions : A. numidicus Grube, Braem's varieties of A. numidicus and A. sud- 
anicus (exc. var. braueri which is doubtfully here for it refers to Brauer's description 
of A. sudanicus), A. somalicus Wedenissow, Sars' three species (1899), A. granarius 
Sars and the nom. nov. asiaticus Gurney, Ueno's var. sinensis and A. sinensis. I 
have seen the types of A. granarius Lucas, A. bottegoi, Bouvier, Apus ovamboensis 
(= cancriformis sens Barnard), and of Tiwari's two species. A. mavliensis Tiwari, 
in spite of its odd appearance is clearly only a young form of this species. A. 
bottegoi Prato is insufficiently described and may belong here or to the previous 


1846. Apus longicaudatus LeConte. 

1852. Apus domingensis Baird. 

1 87 1. Apus aequalis Packard. 

1871. Apus lucasanus Packard. 

1871. Apus newberryi Packard. 

1907. Apus frenzeli Thiele. 

1916. Lepidurus patagonicus, Bruch. (non Berg, 1900). 

1944. Triops pampaneus Ringuelet. 

1947. Apus biggsi Rosenberg. 

1947. Apus oryzaphagus Rosenberg. 

1952. Apus longicaudatus, Linder. 

Range. Western North America, south of 50 N, through Central to South 
America, where only very scattered records (Thiele, 1907 ; Bruch, 1916 ; Ringuelet, 
1944 ; Linder, 1952). W. Indies, Galapagos Islands, Hawaii, Japan and New Cale- 

Habitat. Temporary fresh waters, rice fields in California and Japan. 

Description. As variable as the preceding species, but not so many short bodied 
forms seen. 

Head. — Dorsal organ usually triangular, round in short bodied forms, and anterior 
margin between the eyes. Second maxilla absent in all the specimens seen. 


Carapace. — Text-fig. 2G., h). Shape varies from oval to round with segment 
number. Terminal spine of carina very small when present, but generally 
absent. Carina finely denticulate along whole length in some specimens. Sulcus 
tends to be broad and shallow, spines variable in size, 24-60. 

Body.— Segments : $?, 35-43 ; $S, 35-44- Apodous segments : $?, 5-12 ; <?<?, 
10-13 in material examined. Apodous segments with varying number of super- 
numerary spines on ventral surface. 

Telson. — (Text-fig. 5 D 2 -) Medians large, 1-3, in a single row in the midline. Furcal 
spines smaller than those of T. cancriformis. Posterior marginals very large, set 
in the adult well forward of the margin, often pointing vertically. This arrangement 
of marginals unique and all specimens of this species have it. 

Appendages. — 54-66. (Linder's data mainly) : endites of first thoracic appendage 
variable with segment number. 

Sexual dimorphism. — Well marked in specimens with high segment numbers, 
males then having furcal scales. 

Larvae. — Metanauplius, indistinguishable from that of T. granarius. 

Reproduction. — Bisexual and hermaphrodite, the latter only in California and 
Pacific region populations. 

Fig. 16. Telson of Triops longicaudatus intermedins, (pm = posterior marginals). 

Geographical Races 

1. Triops longicaudatus longicaudatus (LeConte) 

Range. That of species with exception of New Caledonia. 

Description. With the characters described above. There may be a difference 
in the shape of the posterior margin of the telson on the forms from Galapagos and 
Hawaii as against continental American forms ; in the former the posterior margin 
is quite straight with no trace of an emargination in many specimens. This may 
well be evidence for a sub-species inhabiting these islands. Chromosome number 
2W = 8. 

2. Triops longicaudatus intermedins subsp. n. 

Range. New Caledonia. Two samples in MNHNP labelled " 93-1887 " and 


" coll. Simon 25-96 " respectively are of this form and no others from the island are 
known to me. These specimens, all females, form the paratypes of the new sub- 
species. The name suggests that in their telson armature they are intermediate in 
form between their nominate race and the rest of the genus. 

Description. Second maxilla absent, segments 39, apodous 8-9. Dorsal organ 
small, triangular to round. Carapace oval, terminal spine of carina very reduced, 
sulcus spines long, 24-30. Few supernumerary spines on apodous segments ventrally. 
Telson : medians large, 2-3, in mid-line, posterior marginals much smaller than in 
nominate race but well forward of the margin (Text-fig. 16). This is the most char- 
acteristic feature. 


Apus obtusus James, 1823 and Apus guildingi Thompson, 1834 are clearly of this 
species on geographical grounds, but neither description is detailed enough to confirm 
this. Linder has shown that Packard's and Rosenberg's species are synonyms of 
this species. The diagrams of A. frenzeli and Triops pampaneus both show the 
longicaudatus-type telson, and Bruch's record of Lepidurus patagonicus is accom- 
panied by a figure which certainly shows a Triops, and probably T. longicaudatus , 
though the telson is not very clear. 


1896. Apus australiensis Spencer and Hall. 

1905. Apus sakalavus Nobili. 

1907. Apus madagassicus Thiele. 

1 9 1 1 . Triops gracilis Wolf. 

191 1. Triops strenuus Wolf. 

Range. The drier regions of Australia, where it may be locally very common. 

Habitat. Temporary fresh water, sometimes (Wolf, 191 1 ; Main, 1953) in saline 
or alkaline pools and lakes. 

Description. Includes extremes of long and short bodied forms ; from Australia 
several samples have been seen with aberrations (not damage) to limbs and lobes 
of limbs ; both females in a sample from Ayer's Rock have reduced nth thoracic 
appendages so that these are but a stump with a duct running through it on one side 
of each specimen. Several samples from W. Australia have endites 5 and 6 reduced 
to rudiments on one side of the thoracic series. 

Head. — Dorsal organ triangular, ovoid or " wide ", often set with the anterior 
margin well behind the eyes (Text-fig. 4B). Second maxilla absent in all specimens 

Carapace. — Shape variable, terminal carinal spine usually absent and carina 
frequently denticulate along the whole length. Sulcus round, often small, spines 
variable in size, and numerous, 28-62. 

Body.— Segments : ??, 35-43 ; $$, 36-44- Apodous: $$, 5-12; $$, 9-13. 
Apodous segments with varying number of supernumerary spines ventrally. 


Telson. — Medians small, scattered, fewer than in T. granarius, often very few or 
even absent (Text-fig. 6b) ; furcal spines small, posterior marginals small, squat 
and marginal in the adult. Furca varies with the segment number. 

Appendages. — 48-66, but only few specimens counted. Endites variable with 
segment number. 

Sexual dimorphism. — As in previous two species. 

Larvae. — Metanauplius, indistinguishable from that of T. granarius. 

Reproduction. — Bisexual. 

Geographical Races 

1. Triops austr alien sis australiensis (Spencer and Hall) 

Range. Continental Australia. 

Description. With the characters described above. Chromosome number 
2n = 10. 

2. Triops australiensis sakalavus (Nobili) 

Range. Madagascar. 

Description. Very similar to the nominate race, but median spines are rather 
more numerous in the specimens I have seen in MNHNP, which otherwise are clearly 
of this species, and not of T. granarius ; the specimens have no second maxillae, 
the carinae are denticulate, and the median spines are relatively sparse compared 
with T. granarius. 


Wolf's species are clearly Triops australiensis, showing the typical telson pattern 
and Main has already (1953) suggested uniting them with it. T. madagassicus and 
T. sakalavus are placed here on the assumption that only one species will occur in 
Madagascar, since sympatric species are the exception in the Notostraca, but future 
work will be needed to confirm this arrangement. 

Genus LEPIDURUS Leach 

1756. Apus Schaeffer (in part). 

1758. Monoculus Linn, (in part). 

1 8 19. Lepidurus Leach. 

1924. Bilobus Sidorov. 

Distinguished at once from Triops by the presence of a supra-anal plate. In 
general, also, the segment number is lower, the carapace is longer and more com- 
pressed laterally, the sexual dimorphism is weaker than in the other genus. 

Bilobus Sidorov was erected on the erroneous assumption that median incision 
of the posterior margin of the supra-anal plate was important systematically (see 
p. 22, and Linder, 1952). 

ZOOL. 3, 1. 4 



1756. Apus cancrifovmis Schaeffer (in part). 

1758. Monoculus apus Linn, (in part). 

1 801. Apus pro ductus Bosc. 

1 81 9. Lepidurus productus Leach. 

1850. Lepidurus viridis Baird. 

1866. Lepidurus angasi Baird. 

1873. Lepidurus lubbocki Brauer. 

1875. Lepidurus couessii Packard. 

1877. Lepidurus macrourus Lilljeborg. 

1879. Lepidurus viridulus Tate. 

1879. Lepidurus kirki Thompson. 

1879. Lepidurus compressus Thompson. 

1886. Lepidurus packardi Simon. 

1893. Apus extensus Braem. 

1900. Lepidurus patagonicus Berg. 

1909. Lepidurus apus Keilhack. 

191 1. Lepidurus hatcheri Ortmann. 

1921. Lepidurus barcaeus Ghigi. 

1952. Lepidurus couessii, Linder. 

Range. Europe (excluding Britain), North Africa, Palestine, Asia Minor, 
Russia ; North and South America ; New Zealand and Australia. 

Habitat. Temporary fresh waters, but perhaps less restricted to those which 
regularly dry out than, for instance, Triops cancriformis. Occurs in alkaline pools 
in North America. 

Description. This species has the largest range of any known Notostracan 
and shows very little variation over the whole area. 

Head. — Dorsal organ round or oval (Text-fig. 4E, f), anterior margin between the 
eyes with the possible exception of the specimens seen by Ortmann (1911) from Pata- 
gonia. Second maxilla present in all the specimens seen by me. 

Carapace. — Oval, fairly compressed laterally but not so markedly as in next 
species. Terminal spine of carina only rarely absent. Sulcus spines usually long, 
rarely [packardi) reduced. Normally 35-50 in number. 

Body. — Segments, 26-29 > Apodous, 4-6. No supernumerary spines on apodous 

Telson. — Supra-anal plate relatively longer than in next species, the ratio carapace/ 
supra-anal plate being about 4-6 in adults. The marginal spines of the plate are 
small and numerous in all except the South American forms. Central spines 
4-100 or more, the higher numbers being borne on a distinct keel (Text-fig. 7 a) . 

Appendages. — 35-48 ; endites of first thoracic appendages reach to the end of 
the carapace in some, in others little longer than L. arcticus. 

Sexual dimorphism. — Males occur, and the supra-anal plate is longer and more 
spatulate in these ; otherwise the dimorphism is weak. 

Larvae. — Metanauplius in the European and New Zealand forms and probably in 
all. May differ from that of Triops by presence of rudiments of paired eyes in the 
first instar (Bernard, 1892 : 158). 

Reproduction. — Bisexual and hermaphrodite. 

a review of the notostraca 51 

Geographical Races 
1. Lepidurus apus apus (Linn.) 

Range. Europe (exc. range of next ssp.), Asia, North America (exc. California). 

Description. Segments, 26-28 ; apodous, 4-5 ; supra-anal plate with 20-100 
spines generally borne on a keel ; marginal spines of plate fine and numerous. 
Bisexual and hermaphrodite. 

2. Lepidurus apus lubbocki Brauer 

Range. N. Africa, Palestine, Syria, Italy, Sicily. 

Description. Segments, 27-29 ; apodous, 5-6 ; supra-anal plate (Text-fig. 7B) 
with fewer central and marginal spines than above ssp (3-20 centrals) and keel less 
prominent. Specimens from eastern part of the range tend to have the fewest 
central spines ; endites relatively longer than nominate race. Bisexual. Chromo- 
some number, 2n = 12 (Goldschmidt) . 

3. Lepidurus apus packardi Simon 
Range. California. 

Description. Segments and supra-anal plate similar to sub-species 2. Sulcus 
differs from that of all other forms by having very many small spines forming a 
granulated margin. Bisexual. 

4. Lepidurus apus patagonicus Berg 

Range. South America (Chubut Territory, Patagonia). 

Description. Segments, 29 ; apodous, 5. Marginal spines of supra-anal plate 
coarser and fewer than previous forms, central spines few. Sulcus spines as typical 
race. Bisexual. 

5. Lepidurus apus viridis Baird 

Range. New Zealand, Tasmania, and coastal or better watered regions of 

Description. — Very similar to nominate race. Segments, 27-28 ; apodous, 4-5. 
Generally with low number of central spines on the plate (5-10), but one specimen 
from Tasmania had more than 20 ; average is much lower than in nominate race, 
however. I have seen no males. 


I have examined the following types, and am satisfied that they are correctly 
placed here : L. packardi, L. extensus, L. barcaeus, L. angasi, L. cotnpressus, L. kirki, 
L. viridulus. The rest are included on the basis of the original descriptions or on 
material previously determined. 

Lepidurus lemmoni (Holmes, 1894). Holmes' description does not enable deter- 
mination of his specimens with any here recognized, yet contains nothing to indicate 


good differences from them ; the types are lost (Linder, 1952), and therefore it is 
advisable to abandon the species. 


1793. Monoculus arcticus Pallas. 

1883. Lepidurus glacialis Packard. 

1892. Lepidurus spitzbergensis Bernard. 

1893. Apus productus var. glacialis Braem. 
1896. Lepidurus glacialis, Sars. 

1927. Lepidurus ussuriensis Sidorov. 

Range. Circum-polar Arctic regions ; Aleutians, North America ; Alaska to 
Labrador, Greenland, Iceland, Bear Island, Spitzbergen, Northern Palaearctic ; 
Scandinavia to Siberia. 

Habitat. Temporary fresh-water pools, the streams connecting pool systems, 
and large lakes which are permanent. A reservoir in the Norwegian mountains 
(Somme, 1934). 

Description. Head. — Dorsal organ oval, sometimes very elongated, anterior 
margin just between the eyes (Text-fig. 4G). Second maxilla present. 

Carapace. — Oval, laterally very compressed. Terminal carina spine present, long. 
Rest of carina smooth. 

Body. — Segments, 26-28 ; apodous, 4-5. Apodous without supernumerary spines 
on ventral surface. 

Appendages. — 41-46, but variation probably greater. Endites of the first thoracic 
appendage very short, scarcely reaching beyond margin of carapace (Text-fig. ioa). 

Telson. — Supra-anal plate very small ; carapace /plate = about 12 in adults 
of 20 mm. carapace length. Median spines very sparse (0-5) and marginals few 
and coarse (Text-fig. 7c). 

Sexual dimorphism. — Males rare, but known to have longer and more spatulate 
supra-anal plates. 

Larvae. — Post-metanauplius (Poulsen, 1940, and my cultures), equivalent to 
about instar 3 of Triops larvae (Text-fig. 13). 

Reproduction. — Bisexual and hermaphrodite. Males known from Bear Island 
in very low proportion of the population. (Somme, 1934)- 

Geographical Races 

None could be recognized in the material available, nor could Linder (1952) find 
any subdivisions of his material. Lepidurus ussuriensis Sidorov appears to be of this 
species, for the endites are shorter than is usual in other species, and the supra-anal 
plate is small (carapace /plate = about 9) and typical of this species in its armature. 
However, the endites are projecting more beyond the carapace than is usual in 
L. arcticus and may indicate a difference between this East Siberian form and the 
typical Arctic forms. The weakness of the carina and the elongate form of the dorsal 
organ confirm the placing of this species in L. arcticus. 



1883. Lepidurus bilobatus Packard. 
1952. Lepidurus bilobatus, Linder. 

Range. North America ; Utah, Colorado, probably Arizona (Linder, 1952). 

Habitat. Not known, but occurs in arid areas so presumably in temporary 
pools and lakes. 

Description. Known only from Packard's description, and two further specimens 
ascribed to the species by Linder. 

Head. — Dorsal organ round or oval, anterior margin set between the eyes. 

Carapace. — Arrangement of sulcus spines normal. 

Body. — Segments, 33 ; apodous, 6. 

Telson. — Supra-anal plate with numerous small marginals, 4-6 centrals. 

Appendages. — 60 ; endites of first thoracic appendage as in Lepidurus apus. 

Sexual dimorphism. — Males unknown. 

Larvae, Reproduction. — Unknown. 


1952. Lepidurus lynchi Linder. 

1952. Lepidurus lynchi var. echinatus Linder. 

Types. Holotype (?) and allotype ((J) in Uppsala Museum ; paratypes USNM 
82101 ; var. echinatus. Holotype (?) USNM 82068, allotype ((J) USNM 82069. 

Range. North America ; Washington, Nevada, and Oregon. 

Habitat. Apparently in lakes (Linder, 1952 . . . North end of Goose Lake in 
water two feet deep, muddy), but probably also in temporary pools. 

Description. Long bodied form with unique carapace armature. 

Head. — Dorsal organ round with the anterior margin set well behind the eye 

Carapace (Text-fig. 3). — Oval. Carina absent except when it bears series of 
unusually large spines. Specimens from first two localities have 0-20 of these carinal 
spines. Some of those from Oregon have series of large spines along posterior part 
of lateral margin in addition to carinal spines and carapace shape may be more round 
in these. But no useful purpose is served by the formal term var. echinatus for these 
latter forms ; L. lynchi is obviously characterized by the possession of large spines 
on carina and margin and there is much variation in the distribution of these spines ; 
even in the specimens from the first two localities the margin of the carapace bears 
larger spines than is usual in Lepidurus. 

Body. — Segments, 31-34 ; apodous, 3-5. 

Telson. — Supra-anal plate about same size as in L. apus, central spines 2-7, 
marginals very large and sparse. 

Appendages.— 60-71. Endites as in L. apus. 

Sexual dimorphism. —Normal for the genus. 

Reproduction. —Bisexual. 



Types. Holotype (<J) and two paratypes in BMNH (1911 .11.8, 23542-4, Norman 
collection). Collected by W. Bateson. 

Range. Russia. Probably collected at Chilik Kul in the Kazak region, where 
Bateson made a collection of other fresh-water entomostraca also in the Norman 

Description. Males only known. Long bodied form in general appearance 
(Text-fig. 2 J). 

Head. — Eyes and dorsal organ similar to those of Lepidurus lynchi. Second 
maxilla present and typical of the genus. 

Carapace (Text-fig. 2j). Oval-round, more rounded than in Lepidurus apus. 
Carina and its terminal spine absent in all specimens seen ; position of carina demar- 
cated only by the light streak of the dorsal blood channel of the carapace which 
follows the line of the carina in normal forms. Sulcus wide, rounded, with small sulcal 

Body. — Segments 33, apodous 8, in all. Apodous ventral marginal spines very 
small and widely separated centrally, no supernumeraries. 

Appendages. — 49-52. Endites of first thoracic appendage (Text-fig. iod) unique 
in known Notostraca ; endites 4 and 5 very short ; much more reduced than in 
Lepidurus arcticus and can scarcely have projected beyond carapace margin in 
life ; endite 5 of this appendage is little more developed than endite 5 of the second 
appendage. Endite 6 of the first appendage is fully developed and claw-like as in 
the subsequent appendages (in all other Notostraca the 6th endite of the first thoracic 
appendage is reduced to a small soft lobe at the base of the 5th endite). 

Telson (Text -fig. 7E-). — Supra- anal plate very similar to that of Lepidurus lynchi, 
central spines few (4), marginals few, coarse. 

Sexual dimorphism. — Females unknown. 

Reproduction. — Bisexual. 

Note. — One of the paratypes has an abnormality of the supra-anal plate, which 
is reduced in size, soft, and lacks its armature. 


Abonyi, A. 1926. Males of the Apus cancriformis Sch. described on the basis of specimens 
collected in the region of L. Balaton. Arch. Balatonicum, 1 : 71-90. 

Baird, W. 1850. Descriptions of several new species of entomostraca. Proc. zool. Soc. 
Lond. 1850 : 254. 

1852. Monograph of the Apodidae. Ibid. 1852 : 1-7. 

1866. Description of two new species of phyllopodous Crustacea. Ibid. 1866 : 122. 

Balfour-Browne, F. 1909. Note on the rediscovery of Apus cancriformis in Britain. Ann. 

Scot. nat. Hist. 1909 : 118. 
Barnard, K. H. 1924. Contributions to our knowledge of the S.W. African fauna. II, 
Crustacea Entomostraca. Ann. S. Afr. Mus. 20 : 213-230. 

1929. A revision of the South African Branchiopoda Phyllopoda. Ibid. 29 : 181-272. 

Bennie, J. 1894. Arctic plants in the old lake deposits of Scotland. Ann. Scot. nat. Hist. 

3 : 46. 


Berg, C. 1900. Datos sobre algunos crustaceos nuevas para la fauna Argentina. Commun. 

Mus. Nac. Buenos Aires, 1 : 223-236. 
Bernard, H. M. 1889. Hermaphroditism in the Apodidae, Nature, Lond. 43 : 343. 

1892. The Apodidae. A morphological study. London, pp. xvii, 316. 71 figs. 

Borradaile, L. A. 1 926. Notes upon Crustacean limbs. Ann. Mag. nat. Hist. 8 : 193. 

Bosch, L. A. G. 1801. Histoire Naturelle des Crustaces. Paris. 

Bouvier, E-L. 1899. Sur un nouvel Apus de la Somalie. Ann. Mus. Stor. nat. Genova, 

39 : 573-577- 
Bowkiewicz, J. 1923. Biologische Beobachtungen iiber das Vorkommen von Apusidae in 

Siberien. Intern. Rev. Hydrobiol. Leipzig 11 : 329-345, 20 figs. 
Braem, F. 1893. Bemerkungen liber die Gattung Apus. Z. wiss. Zool. 56 : 165-187. 
Brauer, F. 1877. Beitrage zur Kentniss der Phyllopoden. S.B. Akad. Wiss. Wien. 75 : 583- 

Bruch, C. 1916. Dos curiosos crustaceos de San Luis. Physis. 2 : 462-464. 
Campan, F. 1929. Contribution a la connaissance des Phyllopodes Notostraces. Bull. 

Soc. zool. France 54 : 95-118, 8 pis. 
Colosi, G. 1920. Contributo alia conoscenza degli Entomostrachi Libici. Monit. zool. Ital. 

31 : 120-124. 

1922. Note sopra alcuni Eufillopodi. Atti. Soc. Ital. Sci. nat. 61 : 287-297. 

Day, F. 1880. On a new entomostracan from Afghanistan. Proc. zool. Soc. Lond. 1880 : 392. 
Decksbach, N. K. 1924. Zur Verbreitung und Biologic der Apusidae in Russland. Russ. 

hydrobiol. Zeits. 3 : 143-155. 
Fiszera, Z. 1885. Materiaux pour la faune des Phyllopodes de la Pologne. Pam. Akad. 

Fizyogr. 5 : 195-201. 
Fox, H. M. 1945. Haemoglobin in blood-sucking parasites. Nature, Lond. 156 : 475. 

1946. Chemical taxonomy. Ibid. 157 : 511. 

1949. On Apus : its rediscovery in Britain, nomenclature and habits. Proc. zool. Soc. 

Lond. 119 : 693-702. 

1955. The effect of oxygen on the concentration of haem in invertebrates. Proc. Roy. 

Soc, B, 143 : 203. 
Fritsch, A. 1866. Ueber das Vorkommen von Apus und Branchipus in Bohmen. Verhandl. 

zool.-bot. Ges. Wien 16 : 557-562. 
Gauthier, H. 1933. Sur un Apus du Sahare centrale. Bull. Soc. Hist. nat. Afr. N. 24 : 87-96. 

1934- Contribution a l'etude de Y Apus cancriformis et de ses variations dans l'Afrique 

du Nord. Bull. Soc. Sci. nat. Maroc. 14 : 125-139. 
■ 19340. Sur YApus granarius Lucas 1886. Etude du type. Bull. Mus. Hist. nat. 

Paris 6 : 44-46, 2 figs. 
Geikie, A. 1894. Ann. Rep. geol. Surv. Mus. Lond. 1894. 

Geoffroy. 1762. Histoire alregee des insectes qui se trouvent aux environs de Paris, 2 : 658. 
Ghigi, A. 1921. Ricerche sui notostraci di Cirenaica e di altri paesi del mediterraneo. Atti. 
Soc. Ital. Sci. nat. 60 : 161-188. 

1924. Ancora sulla sistematica delle specie mediterranee del genere Triops. Ibid. 63 : 


Goldschmidt, E. 1953- Chromosome numbers and sex mechanism in Euphyllopods. 

Experienta. 9 : 65-69. 
Grasser, J. 1933- Die exkretorischen Organe von Triops {Apus) cancriformis Bosc. Zeit- 

schr. wiss. Zool. 144 : 317-362, 30 figs. 
Grube, E. 1865. Ueber die Gattung Estheria und Limnadia und einen neuen Apus. Arch. 

Naturg. 31 : 203-282. 
Gurney, R. 1921. Fresh water Crustacea collected by Dr. P. A. Buxton in Mesopotamia and 

Persia. /. Bomb. nat. Hist. Soc. 27 : 836-843. 

1923. Notes on some North African and British specimens of Apus cancriformis. Ann. 

Mag. nat. Hist. 11 : 496-502. 

1924. Some notes on the genus Apus. Ibid. 14 : 559-568. 


Guthorl, P. 1934. Die Arthropoden aus dem Carbon und Perm des Saar-nahe-Pfalz- 

gebietes. Abh. Preuss. Geol. Landesamt. 164 : 1-219. 
Hall, R. E. 1953. Observations on the hatching of eggs of Chirocephalus diaphanus Prevost. 

Proc. zool. Soc. Lond. 123 : 95-109. 
Hobson, A. D. and Omer-Cooper, J. 1935. Apus cancriformis in Britain. Nature, Lond. 

135 : 177. 
Holmes, S. J. 1894. Notes on West American Crustacea. Proc. Acad. Sci. Calif. (2) 4 

563-588, 49 figs., pis. 20-21. 
Huxley, J. S. 1942. Evolution, the modern synthesis. London, 1942. 
James, E. 1822. In Long's Expedition to the Rocky Mountains, Philadelphia 1:462 (or 

London, 1823 edition, 2 : 337). 
Keilhack, L. 1909. Zur Nomenklatur der Deutschen Phyllopoden. Zool. Anz. 3 : 177-184. 
Kozubowsky, A. 1857. Ueber den Mannchen von Apus cancriformis. Arch. Naturgesch. 

23 : 312-318. 
Lankester, E. R. 1881. Observations on Apus cancriformis. Quart. J. micr. Sc. 21 : 343- 

Leach, W. E. 1816. Annulosa, Class 1 — Crustacea. Ency. Brit, (supplement to ed. 4-6, 

1 (2) : 404). 
LeConte, J. 1846. A new species of Apus. Amer. J. Sci. Arts, 2 : 274-275. 
Lilljeborg, W. 1877. Synopsis crustaceorum svegicorum ordinis branchipodorum et sub- 

ordinis phyllopodorum. Nova Acta. Soc. Sci. Up sal. 1877 : 1-20. 
Linder, F. 1952. Contributions to the morphology and taxonomy of the Branchiopoda, with 

special reference to the North American species. Proc. U.S. Nat. Mus. 102 : 1-69. 
Linnaeus, C. 1758. Systema Naturae ed. 10. Stockholm. 

Longhurst, A. R. 1954. Reproduction in Notostraca (Crustacea). Nature, Lond. 173 : 781. 
Lundblad, O. 1920. Vergleichende Studien iiber die Nahrungsaufnahme einiger Schwedischeu 

Phyllopoden, nebst synomomischen, morphologischen und biologischen Bemerkungen, 

Ark. Zool. 13 : 1-114. 
Main, A. R. 1953. Sex ratio and variation in Apus australiensis S. and H. W. Aust. Nat. 

4 : 34-39- 
Margalef, R. 1953. Los crustaceos de las aquas continentales ibericas. Biologica de las 

aquas continentales (10) : 1-243. Madrid. 
Mathias, P. 1937. Biologie des Crustacees phyllopodes. Paris, 1937. 
Mayr, E. 1942. Systematics and the origin of species. New York. 1942. 
Moore, E. S. 1893. Some points in the origin of the reproductive elements in Apus and 

Branchipus. Quart. J. Micr. Soc. 35 : 259-284, 2 pis. 
Nobili, G. 1905. Descrizione di un nuovo Apus di Madagascar. Boll. Mus. Torino. 20 (513) : 

Ortmann, A. E. 191 1. Crustacea of Southern Patagonia. Rep. Princetown Univ. Exp. 

Patagonia 1 896-1 899, 3 : 635-667. 
Packard, A. S. 1875. New Phyllopod Crustaceans. Amer. Nat. 9 : 31 1-3 12. 
1883. A monograph of the Phyllopod Crustacea of N. America. U.S. geol. geog. Surv. 

Terr. Rep. 12 : 295-516. 
Pallas, P. S. 1793. Voyages de M. P. S. Pallas . . . , 4 : 1-722. 
Peres, J-M. 1939. Contributions a la connaisance des Notostracees du Maroc. Bull. Soc. Sci. 

nat. Maroc. 19 : 23-32. 
Poulsen, E. M. 1940. Biological remarks on Lepidurus arcticus in East Greenland. Medd. 

Gronland, 131 : 1-50. 
Prato, del, A. 1886. I crostacei della collezione Eritrea Bottego. Atti. Soc. Ital. Sci. nat. 

36 : 1-186. 
Regnard, P. & Blanchard, R. 1883. Note sur la presence de hemoglobine dans le sang des 

Crustacees branchiopodes. C.R. Soc. Biol. Paris, 5 : 197. 
Richters, F. 1886. Ueber zwei Afrikanische ^^s-Arten. Ber. Senckenb. Naturf. Ges. 

1886 : 31. 


Ringuelet, R. 1944. Triops pampaneus nueva especie de Branquipodo Notostraco. Notas 

Mus. La. Plata. 9 : 179-190. 
Rosenberg, L. E. 1947. New species of Apus from California. Trans. Amer. micr. Soc. 

66 : 70-73. 
Sars, G. O. 1899. Additional notes on South African Phyllopoda. Arch. math. nat. Christiana, 

21 : 1-26. 

1901. On the Crustacean fauna of Central Asia. Ann. Mus. St. Petersburg, 6 : 130-164. 

Schaeffer, J. C. 1 756. Der krebsartige Kieffenfuss mit kurzen und langen Schwanzeklappe, 

Abhandlung von Insecten, Regensburg, 1756. 
Schlagintweit, H. 1872. Reisen in Indien und Hochasien, Jena, 3 : 217. 
Schmankiewitsch, W. J. 1 875. Ueber das Verhaltniss der Artemia salina zur Artemia 

mulhauseni und dem Genus Branchipus Schaeffer. Z. wiss. Zool. 25 : 103-116. 
Schrank, F. v. P. 1803. Fauna Boica, Landshut, 3 : 25. 
Sellier, R. & Morice, J. 1946. Le dimorphisme morphologique chez Chirocephalus 

diaphanus Prevost. Bull. Soc. Sci. Bretagne, 20 : 52-54. 
Sidorov, S. A. 1927. Lepidurus ussuriensis sp. nov. Russk Hydrobiol. Z. 6 : 151-152. 
Simon, E. 1886. Etude des Crustaces du sous-ordre des Phyllopodes. Ann. Soc. ent. France. 

6 : 393-4 6 °- 
S0MME, S. 1934. Contributions to the knowledge of the Norwegian fish food animals. I — 

Lepidurus arcticus Pallas. Avh. Norske. Vidensk. Akad. 6 : 1-36. 
Spandl, H. 1926. Wiss. Forschungsergebnisse aus dem Donau und des Schwarzen Meeres. 

2 — Suss. Mikrofauna. Arch. Hydrobiol Stuttgart, 16 : 528-643. 
Spencer, B. & Hall, T. S. 1895. Preliminary description of a new species of Apus, Victorian 

Nat. 11 : 161-162. 

1896. Report of the Horn Society Expedition to Central Australia. London and Melbourne, 

2 : 227-248. 

Summerhayes, V. S. & Elton, C. S. 1923. Contributions to the ecology of Spitzbergen and 
Bear Island. /. Ecol. 11 : 214. 

Tate, R. 1879. Description of a new species of phyllopodous Crustacean. Trans. Proc. phil. 
Soc. S. Australia, 1878-9 : 136. 

Thiele, J. 1907. Einige neue Phyllopoden-Arten des Berliner Museum. Sitz. Ges. Naturf. 
Berlin, 9 : 288-290. 

Thompson, G. M. 1879. On the New Zealand Entomostraca. Trans. Proc. N.Z. Inst. 
1878-1879 : 260. 

Thompson, J. V. 1834. Apus guildingi, a new species. Zoological Researches of J. V.Thomp- 
son, 5 : 108. 

Tiwari, K. K. 1952. Indian species of the genus Apus with descriptions of two new species. 
Rec. Ind. Mus. 49 : 197-206. 

Trusheim, F. 1938. Triopsiden aus dem Keuper-Frankens. Paleont. Z. 19 : 198-216. 

Ueno, M. 1927. Fresh water Branchiopods of Japan. Mem. Coll. Sci. Kyoto. 2 (5) : 259-311. 

1935- Crustacea of Jehol (Phyllopoda). Rep. Sci. Exped. Manchoukuo. Tokyo Section 

5, Div. 1 (2), art. 6 : 1-16. 

1940. Phyllopod Crustacea of Manchoukuo. Bull, biogeog. Soc. Tokyo. 10 : 87-102. 

Wedessinow. T. 1895. Di alcuni Crostacei raccolti nel paese dei Somali dalT ing. Brochetti- 

Robecchi. Bull. Soc. ent. Ital. 26 : 408-424. 
Wolf, E. 191 1. In Michaelsen, W. and Hartmeyer, R. Die Fauna Sud-West Australiens, 

3 (9) : 260. 

Zaddach, E. G. 1 84 1. De Apodis cancriformis — anatome et historia evolutionis. Bonnae. 
Zograf, N. von. 1906. Hermaphroditismus bei dem Mannchen von Apus. Zool. Anz. 
30 : 563-567- 

ZOOL. 3, T. 


2 SEP 1955 




ZOOLOGY Vol. 3 No. 2 

LONDON: 1955 




Pp. 59-148 ; 1 Chart ; 30 Text-figures 



ZOOLOGY Vol. 3 No. 2 

LONDON : 1955 

(NATURAL HISTORY), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical Series. 

Parts appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be compiled 
within one calendar year. 

This paper is Vol. 3, No. 2 of the Zoological series. 


Issued August, 1955 Price One Pound 




i. Introduction .......... 61 

2. Station Details .......... 61 

3. List of Species and Distribution ...... 65 

4. Systematic Section ......... 69 


No large collections of Polychaetes from the Gold Coast have previously been sys- 
tematically studied, though a few records were noted by Augener (1918). The 
present Faunal Survey results from the examination of littoral collections, down to 
a depth of 51 metres, made by Mr. R. Bassindale in 1949-51. Important additional 
material has been collected by Mr. J. B. Buchanan. The author is grateful to these 
gentlemen for the privilege of studying these collections, through which the known 
geographical distribution of many species is extended. Altogether ninety-five 
records of Polychaetes from the Gold Coast are here reported, including two new to 

In the Systematic Section a key to all families, keys to genera and species where 
necessary and a glossary of terms used in the descriptions have been included. All 
previous records from the area have been noted although they may refer to species 
not examined here. References to species have been restricted, as far as possible, 
to standard monographic studies on the group ; the most important are Augener, 
1918, and Fauvel, 1923 and 1927a. 

Dr. A. B. Hastings kindly identified the Polyzoan specimens with which some 
Serpulidae are associated and the author also wishes to acknowledge the technical 
assistance he has received from Miss A. C. Edwards, of the British Museum (Nat. 
Hist.) staff, particularly in the drawing of figures and charts. 


Littoral collections were made at the following places ; the details have been 
provided by Mr. R. Bassindale. 

I. Accra (5 30' N., o° 15' W.) (a) Christiansborg. The reef at Christiansborg 
lies at the foot of the Governor's Castle (2 miles east of Accra lighthouse) 

zool. 3, 2. 6 



and consists of native rock with small deep pools mainly at the lowest 
levels. There is sand and just a little exposed rock above mid-tide level. 
(b) Chorkor. A seine net was operated from the sand beach here, two 
miles west of Accra lighthouse. 
II. Tenpobo and Pram Pram, 24 and 23 miles, respectively, east of Accra. The 
coast is of sand and the upper three-quarters of the beach is a steep sand 
slope. Near the headlands of a series of shallow bays there are low level 
rocks and boulder reefs jutting out to sea. These reefs occupy the lowest 
foot of spring tide range except for native rock, which may rise one or 
two feet above this level. Maximum tidal range is six feet, and neap 
tides do not expose the levels from 0-1 foot above chart datum (low tide 
level of ordinary spring tides) . At Tenpobo the reefs are protected from 
low level wave wash by a line of high rocks to seaward. There are many 
small boulders and stones on the reef ; gravel and sand packing between 
the stones is common. 

III. Winneba, 32 miles west of Accra. The beach here is of native rock from below 

low tide level to above high tide level. 

IV. Apam, 39 miles west of Accra. The shore is of rock and boulders set in sand 

with rock pools. Extending to the east is a shallow sand bay with sandy 
mud in the shelter of the Apam headland. 
V. Sekondi, 113 miles west of Accra. Collections here were made from the hori- 
zontal surface of a rock plateau at or above high tide level. The waves 
wash over the plateau at most states of the tide and so replenish the pools. 
There is probably no period of complete submergence of the plateau. 
VI. Dixcove, 131 miles west of Accra. The bay at Dixcove is almost square, the 
sides being about 400 yards long. The innermost side of the bay has a 
sandy beach with large boulders and native rock at low tide level — 
the rocks rising to mid-tide level or higher. Collections were also made 
on the western side of the bay which consists of small boulders and 
stones, with sand, mud and gravel closer in. 
VII. Princes Town, 142 miles west of Accra. A solid rocky headland very much 
wave-washed, with a few boulders round the headland to the east. 
VIII. Axiwi, 154 miles west of Accra : (a) The Hospital Reef consists of a narrow 
belt of rock, boulders and stones jutting out to sea. Collections here 
were made from the exposed side. 

(b) The Lighthouse Reef juts out in a similar way but is protected by 
two islands, one half way along and one at the seaward end. In-shore 
the reef is at low level and has stones with gravel, sand and mud. Further 
out the stones increase in size until very large boulders occur on the 
exposed side. Collections here were made mainly from the sheltered 

Off-shore collections were made by Mr. Bassindale entirely within the area off 
Accra shown on the chart (p. 63). The following lists give further details of the 
stations. Several collections were made by Mr. Bassindale at stations which are not 



6 4 


numbered and they are referred to in the text by reference to points on the coast. 
Mr. Buchanan made part of his survey within the area indicated on the chart between 
3*6 and n-o metres ; other stations at which he made collections have also been 
referred to as the Buchanan Survey and their positions have been indicated in the 
same way. 

List of Collecting Stations in the Sea off Accra 

All dredge hauls of 10 minute duration and Agassiz Trawl hauls of 15 minute 
duration except where stated. 


Depth in 




Instrument used 




Large dredge 




» >» 




9 min 
























5 min 




8 min 
















30 min 








Small dredge 15 min 



Large dredge 



Small dredge 







> >> 



• >> 



• > > 
























, , 


x 4 






















Agassiz Trawl 










List of Collecting Stations in the Sea off Accra — cont. 


Depth in 




Instrument used 




Agassiz Trawl 










„ 30 mm 













20 mm 







No details 




Agassiz Trawl 





12. iv. 51 









14. iv. 51 














Approximate distance covered in normal 10-minute haul with large dredge was 700 metres, 
in normal 15 -minute haul with Agassiz Trawl, 400 metres. (More in shallow and less in deep 


X = Collected in the present survey. 

o = Recorded by Augener (1918), not collected in the present survey. 





Aphrodite alta 
Hermione hystrix 
Panthalis bicolor 
Euthalenessa insignis 
Eusigalion vazensis 
Sthenelais boa . 

Leanira japonica 
,, yhleni 








t— 1 





















List of Species and Distribution — cont. 

X = Collected in the present survey. 

o = Recorded by Augener (1918), not collected in the present survey. 



Scalisetosus pellucidus 
Acholoe astericola 
Pareulepis geayi 
Lepidonotus hupferi . 
Harmothoe goreensis 

5 g 

O o3 













Bhawania goodei 

Chloeia viridis . 

Hermodice carunculata var didymo- 

branchiata . 
Eurythoe complanata 



Phyllodoce africana . 
oculata . 
Eteone picta 


Leocrates claparedii . 

Trypanosyllis prampramensis 
Syllis gracilis . 
„ hyalina . 
,, cornuta . 
Opisthosyllis brunnea 
Pionosyllis ehlersiaeformis 
Exogene gemmifera . 

Nereis falsa 
Neanthes succinea 
Perinereis melanocephala . 
Pseudonereis gallapagensis 



. . X 

x . . X 

. . . . X 



List of Species and Distribution — cont. 

X = Collected in the present survey. 

o = Recorded by Augener (19 18), not collected in the present survey. 




Nephthys hombergii . 
Aglaophamus lyrochaetus 


Glycera convoluta 
Goniada multidentata 
Goniadopsis incerta . 
Ophioglycera archeri sp. n. 


Eunice antennata 
,, vittata . 
rubra . 
Marphysa dartevellei 
Lysidice ninetta 
Diopatra musseraensis 

,, neapolitana 
Onuphis eremita 
Lumbrineris impatiens 
,, albifrons 

Holla parthenopeia . 

Orbinia foetida var linguistica 
Scoloplos madagascarensis 
,, dubia, sp. n. 

Prionospio pinnata . 


Audouinia tentaculata 
,, filigera . 

punctata . 
Cirratulus filiformis . 









u , — 

B 3 


x x 


n O 

< in 






r— I 









. . X 





. . . . X X X 

X . . X 

X . . 

X . . X 



List of Species and Distribution — cont. 

X = Collected in the present survey 

o = Recorded by Augener (1918), not collected in the present survey. 



StylaHoides scutigeroides . 
„ arenosus 

„ tropicus 


Armandia intermedia 
Polyophthalmus pictus 


Maldane sarsi 


Owenia fusiformis . 


Sabellaria eupomatoides 

spinulosa var intoshi 
,, spinulosa var alcocki 

Sternaspis scutata var africana 


Phyllamphicteis collaribranchis 
Pterolysippe bipennata 
Isolda whydahensis . 

Pista grubei 
Terebellides stroemi 
Streblosoma persica 
Loimia medusa 
Amaea accraensis 

Laonome puncturata 
Dasychone lucullana 
Potamilla casamancensis 











t— ( 






Augener (1918), from Saltpond. 




X X 

x . . X 







X . . X 






List of Species and Distribution— cont. 

X = Collected in the present survey 

o = Recorded by Augener (1918), not collected in the present survey. 


„ , — .> . — . 



Salmacina incrustans 
Hydroides norvegica 
spinosus . 
,, arnoldi 

Apomatus similis 
Serpula vermicularis 
Vermiliopsis multicristata 

Spirorbis sp. . 

ft U 

6 p., 










. X 

. X 

. X 



X X 
















x . . .. 

. . . . X 

Augener (191 8) from the Cape Coast Castle. 


(a) Key to Families of Polychaeta 

The following key has been adapted from Fauvel (1923 and 1927a). It includes 
families not collected in the present survey and these have been marked with an 

1. Some parapodia bearing elytra, the rest with dorsal cirri . . . Aphroditidae. 
Parapodia without elytra . . . . . . . . . . 2. 

2. A fan-shaped group of broad, flattened chaetae (paleae) on all segments 

No such groups of chaetae .......... 3. 

3. Prostomium not distinct ; when present pedal cirri globular . . . . 4. 
Prostomium distinct ......... .6. 

4. Parapodia biramous and achaetous ; prostomium fused with the two succeeding 

segments and bearing 2 or 4 tentacular cirri containing aciculae ; pedalcirri absent. 
Pelagic ........... Tomopteridae.* 

Parapodia uniramous, with chaetae and with globular cirri ..... 5. 

5. Pharynx armed .......... Pisionidae.* 

Pharynx unarmed ......... Sphaerodoridae.* 

6. Body not normally divided into distinct regions ; all segments alike except those 

near the mouth and the pygidium. Pharynx protrusible and normally armed 
with jaws. This group includes those families of the Polychaeta Errantia of 
Fauvel (1923) not given above ......... 7. 


Body often divided into distinct regions and adapted to either a tubicolous or 
burrowing life, the head is, therefore, either greatly modified, with a wreath of 
tentacles or gills surrounding the mouth, or almost completely divested of cephalic 
appendages. Parapodia much modified in the tubicolous forms in which the 
neuropodia are often in the form of pinnules (tori) bearing hooks or uncini. 
Without jaws. This group includes the Polychaeta Sedentaria of Fauvel, 

1927a 15- 

7. Prostomium conical, without antennae or palps ; dorsal and ventral cirri foliaceous ; 

chaetae rare, when present simple, acicular. Pelagic . . Typhloscolecidae.* 

Prostomium with tentacles, usually with palps . . . . . . . 8. 

8. Prostomium small with three antennae ; pharynx unarmed ; a caruncle almost 

always present. Chaetae simple, calcareous in some genera . Amphinomidae. 

Prostomium prominent ; without a caruncle . . . . . . . 9. 

9. Prostomium long, conical, annulated, with four short terminal antennae Glyceridae. 
Prostomium not so ........... 10. 

10. Proboscis without chitinous teeth . . . . . . . . .11. 

Proboscis with chitinous teeth . . . . . . . . . .14. 

11. Parapodia normally uniramous, cirri usually foliaceous . . . . .12. 
Parapodia biramous or very much reduced . . . . . . 13. 

12. Prostomium with two very large globular eyes. Pelagic . . . Alciopidae.* 
Prostomium with two small eyes ....... Phyllodocidae. 

13. Parapodia biramous, with inter-ramal cirri (branchiae) . . . Nephthydidae. 
Parapodia biramous, or very much reduced, without branchiae . . Hesionidae. 

14. Proboscis with one pair of heavy curved jaws distally and normally with paragnaths 

on its dorsal and ventral surfaces ........ Nereidae. 

Proboscis with a complex armature — a pair of ventral pieces and two, or frequently 

more, dorsal pieces ......... Eunicidae. 

Proboscis with one large tooth, or crown of teeth, or both .... Syllidae. 

15. Body clearly divided into distinct regions . . . . . . . 23. 

Body not clearly divided into distinct regions . . . . . . 16. 

16. Body very much swollen, few segments, filiform anal branchiae. A large ventral 

shield bordered with chaetae ....... Sternaspidae. 

Body not swollen, numerous segments, without anal branchiae. No ventral 

shield ............. ij. 

17. Palps elongate ............ 18. 

Without palps ............ 22. 

18. Elongate palps (tentacular cirri) on the body, not on the prostomium . Cirratulidae. 
Elongate palps on the prostomium . . . . . . . . ig. 

19. A cephalic cage formed by the chaetae of the first chaetigers . Flabelligeridae. 
Without a cephalic cage . . . . . . . . . . .20. 

20. Without branchiae. Prostomium oval, spoon shaped. Palps with sucker-like 

papillae ........... Magelonidae.* 

With branchiae ............ 21. 

21. Parapodial lamellae erect, dorsal. Branchiae cirriform. Hooded hooks and 

capillary chaetae .......... Spionidae. 

Anterior parapodial lamellae frilled or flask-shaped. Branchiae filliform. Chaetae 

of many kinds, plumed, spiny, barbed and acicular, etc. . . . Disomidae.* 

22. One median antenna. Dorsal foliaceous branchiae. Hooded hooks and capillary 

chaetae ........... Paraonidae.* 

Two antennae or none. Capillary and forked chaetae, no hooded hooks 


Prostomium blunt, without appendages or with a crown of lobes. No branchiae. 

Ventral tori with rows of uncini. Sandy tube ..... Oweniidae. 

Prostomium without appendages, with a keel or rimmed cephalic plate . Maldanidae. 













A terminal branchial tuft with numerous filaments. Thoracic uncini dorsal 
abdominal uncini ventral ........ 

No terminal branchial tuft ........ 

Paleae forming an operculum to close the tube ..... 

No opercular paleae ......... 

Prostomium without processes. Branchiae on many segments, but not on the 
prostomium . . . . . . 

Prostomium with processes, either branchiae or tentacular filaments 

With uncini ........... 

Without uncini .......... 

With acicular hooks or capillary chaetae ...... 

With no acicular hooks, only capillary chaetae . 
Prostomium blunt. Non-retractile arborescent 

branchiae in the 




middle body 
region ........... Arenicolidae.* 

Prostomium conical. Branchiae, if present, posterior in position, simple or multifid, 
if multifid retractile ......... Capitellidae.* 

Body divided into 2-3 strikingly dissimilar regions. Prostomium with or without 
2 small tentacles. Two long grooved palps. Anterior region short with uni- 
ramous feet bearing peculiar chaetae in the 4th chaetiger. Posterior notopodia 
erect. Uncini comb-like ....... Chaetopteridae.* 

Body divided into regions which are not strikingly dissimilar. With numerous 

tentacular filaments . . . . . . . . . . 30. 







Tentacular cirri retractile into the mouth. Prostomium distinct . 

Tentacular cirri not retractile into the mouth. Prostomium indistinct 

An operculum of one anterior row of golden paleae 

An operculum of two stalks bearing a crown of paleae 

Tube calcareous, normally with an operculum .... 

Tube membranaceous or mucous, never with an operculum . 

(b) Glossary of Terms 

The following glossary includes only those terms which are not normally to be 
found in a standard text-book of zoology or English Dictionary. 

Acicula : A stout chaeta of one piece, almost always tapering to a blunt point. 

Capillary Chaeta : Simple chaeta, normally long and slender. 

Caruncle : A longitudinal ridge on the dorsal surface of the prostomium and sometimes 

extending to some of the following segments. 
Chaeta : A bristle made largely of chitin but exceptionally some may be composed of calcium 

carbonate (e.g. Eurythoe complanata). Chaetae are either simple (of one piece), or com- 
pound (having two or more articulating pieces). 
Falciger : The terminal article of a compound chaeta when it is short and broad. c.f. 

Heterogomph : The condition when the two lateral branches of the articulating surface of 

the main stem of a compound chaeta are unequal in length. 
Homogomph : The condition when the two lateral branches of the articulating surface of the 

main stem of a compound chaeta are equal in length. 
Limbate : The condition when a simple chaeta is bordered along all or part of its length, e.g. 

Apomatus similis, Text-fig. 30, /. 
Palea : A stout simple chaeta, groups of which appear in the Chrysopetalidac, Amphictenidae, 

Ampharetidae and Sabellariidae. 
Sesquiramous : Condition of a parapodium when the dorsal component (notopodium) is 

reduced to a few chaetae or an acicula. 


Spiniger : The terminal article of a compound chaeta when it is long and slender, c.f. 

Stylode : A dorsal appendage of the branchial filaments in some Sabellidae. 
Uncinus : A short simple chaeta which may be sigmoid, with teeth at the tip, or plate-like 

with rows of teeth along one edge. 

(c) Descriptions of Species 

Keys to genera and species have been compiled only in respect of forms actually 
recorded from the Gold Coast, or where systematic considerations suggest that details 
of related forms will be of help to the student. 

Examples of all species have been deposited in the collections of the B.M. (N.H.), 
Reg. Nos., 1953.3.1.1-1234 ; duplicate material covering the majority of species, 
has been deposited in the Zoology Dept., University College, Achimota, Gold Coast. 

Family Aphroditidae 

Prostomium normally with four sessile, or pedunculate, eyes, one to three antennae 
and two palps inserted ventrally. Protrusible proboscis crowned with papillae 
and with four chitinous jaws (except the Aphroditinae) . A feature of this family 
is the possession of elytra on the dorsal surface of the parapodia. The first pairs 
are always attached to segments 2, 4, 5 and 7, thereafter the arrangement varies, 
posteriorly they may be absent. Parapodia biramous, exceptionally reduced. 

Key to Subfamilies 

1. Body short, fat and oval, jaws absent or rudimentary. Elytra may be covered with 

a thick mat of felt ......... Aphroditinae. 

Body comparatively long and thin, pharynx with jaws. Elytra never covered with 

a mat of felt ............. 2. 

2. Compound chaetae in the neuropodia. Elytra on all segments posteriorly Sigalioninae. 
All chaetae simple. Elytra on every second or third segment, or entirely absent, 

from the posterior part of the body . . . . . . . . . 3. 

3. Elytra on every second segment in the posterior part of the body. Prostomial eyes 

sessile or stalked. Tubicolous in habit ...... Polyodontinae. 

Elytra on every third segment in the posterior part of the body, or entirely absent. 

Prostomial eyes sessile, never stalked. Errantiate in habit . . Polynoinae. 

Subfamily Aphroditinae 

One unpaired antenna, no laterals ; a facial tubercle beneath the antenna. 
Elytra 15 pairs, rarely 20, inserted on segments 2, 4, 5, 7, 9-21, 23, 25, 28, 31, 
alternating with dorsal cirri. 

Key to Genera 

1 . Dorsal surface covered with a thick felt ; dorsal notopodial chaetae ending in blunt 

points or with fine hooks ......... Aphrodite. 

2. No covering of felt ; notopodial chaetae harpoon-shaped on elytrigerous segments 



Genus APHRODITE Linnaeus, 1758 
Fifteen pairs of elytra. Eyes sessile or absent. 

Aphrodite alt a Kinberg, 1856 

Aphrodite alta, Mcintosh, 1924, pp. 5-6, from Cape of Good Hope waters. 

Aphrodite alta, Monro, 1930, pp. 36-38, fig. $a-i, from 64 20' 00" S., 63 01 ' 00" W. 

Localities. Off Accra : Stn. 47 (1) ; Stn. no (1) ; Stn. 133 (1) ; all complete, 
20 mm. long by 10 mm. wide. 

This species is characterized by the lack of eyes, the dorsal notopodial chaetae 
which taper to a fine hooked point, and which appear to lie in the dorsal felt rather 
than to project through it, the thick bearded ends of the neuropodial chaetae and 
the posterior neuropodial chaetae with long alternating teeth. These features 
clearly distinguish the species from A. aculeata which Fauvel (1950) reported from 
Rio de Oro. 

Distribution. This appears to be the first record of A. alta from the W. coast 
of Africa, north of the Cape. Kinberg (1856) reported it from Rio de Janeiro. 

Genus HERMIONE Blainville, 1828 
Four pedunculate eyes. Fifteen pairs of elytra. 

Hermione hystrix (Savigny), 1818 
Hermione hystrix, Fauvel, 1923, pp. 35-36, fig. 11. 

Locality. Off Accra, Stn. 126 (12), all complete, measuring between 20- 
30 mm. long by 10-15 mm. wide. 

There are two pairs of eyes, each pair carried on a peduncle, situated ventral to 
the antenna. Two types of notopodial chaetae are present on elytrigerous segments ; 
one set, stout, curved and ending in a simple point, the others with strong, harpoon- 
shaped points. On cirrigerous segments there are only thin chaetae with simple 
points. Neuropodial chaetae are bifurcate with a short limb and a long curved one 
which may be toothed on anterior and posterior feet. 

Distribution. Mediterranean Sea, Atlantic Ocean and Indo-Pacific regions. 

Subfamily Polyodontinae 

Bilobed prostomium, with four sessile eyes, or two large ommatophores and three 
antennae. Elytra on segments, 2, 4, 5, 7, 9, etc. 

Only one genus of this subfamily has been recorded from the Gold Coast, Pan- 
thalis, but the following Key is included here to illustrate the difference between 
it and Polyodontes. 

1 . Superior neuropodial chaetae elongate hastate, the tip smooth or more or less hirsute 


2. Superior neuropodial chaetae penicillate (Text-fig. 1) . . . . Panthalis. 


Hartman (1939) has drawn attention to the need for revising the diagnostic generic 
characters in this subfamily and this Key has been adopted from her suggestions. 
Thus the presence or absence of branchial lobes is of doubtful value, for many speci- 
mens are incomplete when collected and the methods adopted for preservation fre- 
quently result in the complete retraction of these organs. 

Genus PANTHALIS Kinberg, 1855 
Panthalis bicolor Grube, 1877 

Polyodontes bicolor, Augener, 1918, pp. 119-125, pi. 2, fig. 28 ; pi. 3, fig. 47 ; text-fig. 4. From 

Localities. Off Accra, Stn. 33 (1), anterior fragment 23 mm. in length ; Buchanan 
Survey, in a depth of 3-6 to n -o metres (1), anterior fragment, 10 mm. long ; Buch- 
anan Survey, in a depth of 10 metres off lighthouse (1), anterior fragment, 17 mm. 

The prostomium carries two large anterior ommatophores (eye-stalks) and two 
small posterior eye-spots, situated on either side of the median antenna, which 
projects forward between the ommatophores. Two lateral antennae are ventral 
to the eye-stalks. There are two pairs of tentacular cirri. Spinning glands are 
present in the feet. Neuropodial chaetae are illustrated in Text-fig. 1, (a) superior 
penicillate with the tip frequently carried beyond the bushy portion, (b) median 
stout aristate, (c) inferior serrulate. 

No tubes were found with these specimens. 

Distribution. The species is known only from the tropical West African coast. 

Subfamily Sigalioninae 

Elytra numerous, attached to segments 2, 4, 5, 7, etc., and on each segment from 
the 23rd-29th up to the end of the body. All the genera here reported have three 
antennae ; other genera, however, have one (Pholoe) or two (Sigalion). 

Key to Genera 

1. With a dorsal cirrus on the 3rd chaetiger . . . . . . . . 2. 

Without a dorsal cirrus on the 3rd chaetiger ....... 3. 

2. With the unpaired antenna inserted on the anterior margin of the prostomium 

With the unpaired antenna inserted on the middle of the prostomium, in line with, 

or posterior to, the eyes .......... Eusigalion. 

3. With falcigerous compound chaetae in the neuropodium .... Sthenelais. 
With spinigerous compound chaetae in the neuropodium .... Leanira. 

Genus EUTHALENESSA Darboux, 1899 

The median unpaired antenna, without ceratophore or ctenidia, is on the anterior 
margin of the prostomium. From the fourth chaetiger dorsal ctenidia are present 
on the parapodia ; on the third foot there is a branchial tubercle but no true dorsal 



Fig. i. Panthalis bicolov. Neuropodial chaetae ; (a) superior penicillate ; (b) median 
aristate ; (c) inferior serrulate, x 200. (Specimen from the Buchanan Survey.) 

Euthalenessa insignis Ehlers, 1908 

Euthalenessa insignis Augener, 1918, pp. 108-112, from Apam. 

This species has not been found in the present collections. 

The specimens described by Augener were characterized by the possession of a 
third cirrus on the first chaetiger, and it is doubtful if this character is applicable 
to Ehler's species, but specimens will have to be examined before the true identity 
of Augener's record can be established. 

Genus EUSIGALION, Augener, 1918 

The median antenna is in the middle of the prostomium, between, or posterior to, 
the eyes. This character is here taken to separate Eusigalion from Euthalanessa, 
which are similar in other respects. 

ZOOL. 3, 2. 7 

7 o 


Eusigalion vazensis Augener, 1918 

Eusigalion vazensis Augener, 1918, pp. 113-118, pi. 2, fig. 14 ; pi. 3, figs. 44-46 ; text-fig. 3. 
From Fernand Vaz in the French Congo. 

Locality. Off Accra, Buchanan survey, in 3-6 to n-o metres depth (2), anterior 
pieces, 20-23 mm. long. 

There are about 12 pinnate filaments on the outside edge of each elytron. An- 
tennae are very short, the unpaired median is set back on the dorsal surface of the 
prostomium, between the two pairs of small eyes ; the paired anterior antennae 
project from the front border of the prostomium. Notopodial simple chaetae are 
long, barred and serrated. Neuropodial chaetae are in two groups ; a supra- 
acicula group of simple spinose bristles and long-shafted compound forms with 
numerous joints, and a sub-acicula group, all compound, one set with single long- 
shafted, bifid, terminal articles, the other with numerous articles of which the 
terminal one is also bifid. 

Distribution. Only known from the above records. 

Genus STHENELAIS Kinberg, 1855 

Prostomium with four eyes. The lateral antennae are fused with the first chaetiger. 
Unpaired median antenna with basal ctenidia. Two long palps with ctenidia at the 
base. Branchiae on all parapodia from the fourth. Notopodial chaetae simple ; 
neuropodial chaetae compound f alcigers with sometimes a superior group of simple 

Key to Species 
1 . Elytra with simple fringes and entire borders ...... 

Anterior elytra with bifid fringes ; 
border notched 

S. boa. 
elytra from the 24th chaetiger with outside 

5. limicola. 

Sthenelais boa (Johnston), 1833 
Sthenelais boa, Fauvel, 1923, pp. 110-111, fig. 41, a-l. 

Locality. Tenpobo shore, 6.U.50 (1), 43 mm. long. 

Elytra normally reniform, crossing and overlapping on the back, with minute 
papillae and with simple fringes on the outer border. Notopodial chaetae simple, 
finely spinous. Superior neuropodial chaetae simple, bi-pectinate, strongly spinous, 
the middle group compound f alcigers with single bidentate articles, the inferior group 
also compound with long, pseudo-multiarticulate, bidentate falcigers. 

Distribution. English Channel, Mediterranean Sea, Atlantic Ocean, Indian Ocean. 

Sthenelais limicola (Ehlers), 1864 

Sthenelais limicola, Fauvel, 1923, pp. 113-114, fig. 42, a-g. 

Locality. Off Accra, Buchanan Survey, in 3-6-11-0 metres depth (4), two com- 
plete, 60, 75 mm. in length, two fragments 23 and 56 mm. long. 


The characters noted in the above key serve to separate this species from S. boa ; 
in other characters the species are similar. 
Distribution. North Sea, Mediterranean Sea, Atlantic Ocean. 

Genus LEANIRA Kinberg, 1855 

Members of this genus may, or may not, have eyes ; otherwise they are similar 
to Sthenelais, except as keyed above. 

Key to Species 

1 . Superior neuropodial chaetae simple, with whorls of spikes ; inferior neuropodial 

chaetae compound, caniculate spinigers . . . . . . L. japonica. 

2. All neuropodial chaetae compound, caniculate spinigers . . . L. yhleni. 

Leanira japonica Mcintosh, 1885 

Sthenolepis japonica, Willey, 1905, pp. 259-260, pi. 2, rig. 49. 
Leanira japonica, Fauvel, 1932, pp. 33-34. 

Localities. Off Accra, Stn. 57 (1) ; Stn. 70 (3) ; Stn. 97 (1) ; Stn. 132 (1) ; Stn. 
I 33 ( 2 )- One of the specimens from Stn. 70 is complete, 64 mm. in length. All 
others are fragmentary, up to 15 mm. long. 

Prostomium with four eyes. The dorsal cirrus is replaced in this species by a 
very small tubercle. The elytra, which are fringed, overlap but do not cross, and 
leave a considerable area of the mid-dorsal surface exposed. Notopodial chaetae 
are long and slender, neuropodial compound chaetae have long caniculate terminal 
pieces and a superior group of simple chaetae, with whorls of spikes, on most 

Distribution. Previously reported from Japan, Malaya, Bay of Bengal and the 
Arabian Sea, this appears to be the first record of this species from the Atlantic 

Leanira yhleni Malmgren, 1867 
Leanira yhleni, Fauvel, 1923, pp. 117-118 (no figures). 

Locality. Off Accra, Buchanan Survey, 41*4 metres depth, off Lagoon, (1), 
anterior fragment 25 mm. long. 

This species is distinguishable from L. japonica as noted above in the key, and in 
having the anterior pair of eyes larger than the posterior pair. In L. japonica the 
posterior eyes are larger than the anterior. In both species the anterior eyes are 
near the front border of the prostomium, partly hidden beneath the antennal ctenidia, 
and the posterior pair are at the base of the unpaired antenna. 

Distribution. Bay of Biscay, Mediterranean Sea, Adriatic Sea ; this appears 
to be the first record from the equatorial Atlantic. 

Subfamily Polynoinae 

Elytra 12 or more pairs inserted on segments 2, 4, 5, 7, 9, to 23-26, thence on every 
third segment to the posterior end of the body. All genera here reported have one 
unpaired, median antenna and two paired, lateral antennae. 


Key to Genera 

i. Lateral antennae inserted ventrally ; 15 pairs of elytra . . . . . 2. 

Lateral antennae inserted terminally, continuous with the prostomial peaks . . 3. 

2. Chaetae transparent as crystal, with spinous pouches .... Scalisetosus. 
Chaetae not transparent, without spinous pouches ..... Harmothoe. 

3. Elytra numerous, more than 15 pairs, a large dorsal tubercle in the form of a T on 

the cirrigerous segments ......... Acholoe. 

Elytra 12 pairs ............ 4. 

4. Segment 3 fused with the segments 2 and 4 dorsally ..... Pareulepis. 
Segment 3 not fused dorsally ........ Lepidonotus. 

Genus SCALISETOSUS Mcintosh, 1885 

The 15 pairs of elytra are inserted on segments 2, 4, 5, 7-23, 26-29-32, not covering 
the whole body. Neuropodial chaetae with semilunar cusps are slightly bifid at the 
extremity. Notopodial chaetae curved with blunt spines. 

Scalisetosus pellucidus (Ehlers), 1864 
Scalisetosus pellucidus, Fauvel, 1923, p. 74, fig. 27, a-f. 

Locality. Off Accra, stn. 47 (1), 10 mm. in length. 

Irregular dark brown transverse bands decorate the dorsal surface. Antennae 
and cirri are papillated. The ventral chaetae are not as deeply incised at their distal 
ends and the notopodial chaetae not so clearly notched as normal representatives 
of the species. 

Distribution. Mediterranean Sea, Atlantic Ocean, Bay of Bengal, Malay 

Genus ACHOLOE Claparede, 1870 

Two pairs of tentacular cirri with basal aciculae. Elytra on segments 2, 4, 5, 7-23, 
26, 29, 32, thence almost up to the end of the body. Parapodia with reduced noto- 

Acholoe astericola (Delle Chiaje), 1823 

Acholoe astericola, Fauvel, 1923, pp. 94-95, fig. 36, d-h. 

Localities. Off Accra : From starfish, at 25 metres, and 10 metres (off Petrol 
Buoy), numerous specimens; Stn. 57 (1), among tubes of Diopatra neapolitana ; 
Stn. 67 (1) ; Stn. 80 (1) ; Stn. 89 (1) ; Stn. 104 (1) ; Stn. 123 (1). 

Most of these specimens are fragmentary, the complete ones measure up to 30 mm. 

There are about 45 pairs of elytra. Neuropodial chaetae have a recurved un- 
identate tip with a short spinous region, and notopodial chaetae are short with trans- 
verse rows of pinnules. 

Distribution. English Channel, Mediterranean Sea, Atlantic Ocean. 



Genus PAREULEPIS Darboux, 1899 

Segment 3 fused with segments 2 and 4 dorsally. Elytra on segments 2, 4, 5, 7, 
>, 11-21-24. Notopodial chaetae are slender capillaries and stout curved bristles 
>ent abruptly at their distal ends. Neuropodial aciculae are short, wholly embedded, 
vith extended, plate-like terminations. Neuropodial chaetae include a few small, 
aterally pectinate, bristles, and a bundle of almost straight chaetae. 

Pareulepis geayi (Fauvel), 1918 
lulepis geayi, Fauvel, 1919, PP- 335-339, pi- 15, figs. 17-21 ; pi. 17, figs. 76-79. 

Locality. Off Accra, Buchanan Survey, between 11-13 metres depth, off 
'hristiansborg Castle (2), each about 25 mm. long. 

The arrangement of anterior segments in this species is shown in Text-fig. 2. Thus 
egment 3 is represented laterally by parapodia, but is fused dorsally with segments 





Fig. 2. Pareulepis geayi. Dorsal view of anterior segments to show the fusion of 
segment 3 with segments 2 and 4 dorsally. Elytra and the 2nd and 3rd parapodia on 
the left-hand side have been omitted. 

5 and 4. Elytra are present therefore on segments 2, 4, 5, 7, 9-21-24, after which 
here are ten cirrigerous segments up to the end of the body. The elytra on segment 
*4 extends from segment 23 backwards to 29, so that only a small posterior portion 
>f the worm remains uncovered. There are up to 12 marginal papillae on the outside 
border of each elytron. Dorsal cirri are stout and foliaceous throughout. 

The specimens here described have been referred to this species, and not to P. 
Imbriata Treadwell, with which Hartman (1939) considers P. geayi synonymous, 


because of the position of the unpaired median antenna. Fauvel (1919, p. 338) 
clearly implies that the unpaired antenna is posterior to the median line of the pro- 
stomium. Treadwell (1900, p. 190), however, places it on the anterior prostomial 
margin. Too, Treadwell's specimens had eye-spots ; these are not present in Fauvel's 

Distribution. This species appears to have been recorded only from the Indo- 
Pacific region, though the genus, through P.fimbriata, is also known from the Pacific 
coast of America. 

Genus LEPIDONOTUS Leach, 1816 

Prostomium bilobed, without frontal horns, four eyes. Twelve pairs of elytra 
attached to segments 2, 4, 5, 7-21, 23. 

Lepidonotus hupferi Augener, 1918 
Lepidonotus hupferi Augener, 1918, pp. 133-136, pi. 2, figs. 7-1 1 ; text-fig. 5. 

Localities, (i) Tenpobo shore (10), most complete, between 8-18 mm. in length, 
collected in February, 1950. (2) Off Accra, from wood netted off Chorkor in 14 
metres, 14.V.51 (1), 10 mm. long. (3) Axim, (a) Shore, 13/14.^.49 (1), 12 mm. long ; 
{b) Hospital Reef (4), 2 complete, 4 and 6 mm. long ; 2 parts 4 and 8 mm. long. 

The elytra completely cover the dorsal surface, are fringed on their external 
margins, and covered with very small tubercles, with a broad base and slender stalk. 
Notopodial chaetae are slender and serrated. Neuropodial chaetae are stout with a 
serrulated blade ; some are distally entire, others have a slightly bifid tip. 

Fauvel (1950) has noted that this species appears to replace L. squamatus in the 
tropical zone. 

Distribution. Tropical West Africa, Western Mexico, Panama, N.W. South 
America. Hartman (1939) records this Western Hemisphere distribution from the 
intertidal zone to 22 metres. 

Genus HARMOTHOE Kinberg, 1855 

Prostomium bilobed, with four eyes, often with lateral peaks. Elytra covering 
all the dorsal surface on segments 2, 4, 5, 7-23, 26, 29, 32. 

Harmotho'e goreensis Augener, 1918 
Harmothoe goreensis Augener, 1918, pp. 142-146, pi. 2, figs. 4-6 ; pi. 3, fig. 42 ; text-fig. 7. 

Localities. Off Accra : (a) Stn. 47, (1), 8 mm. long ; (b) from wood netted off 
Chorkor in 14 m., (4) ; (c) 2 miles out from R. Densu, 8.iv.49, 14-6 m., in shingle, 
(4), 2 complete, 6 mm. long ; (d) 2 miles west out from R. Densu, 1 mile off shore, 
2.iii.49, 7-3 m., (1), 4 mm. long. 

The elytra have short fringes on their external margins ; elytron tubercles are 
sac-like in shape with a minute distal pore. Tentacular cirri, dorsal cirri and palps 
are covered with papillae. Neuropodial chaetae are normally bidentate superiorly 


and inferiorly are unindentate. Notopodial chaetae (always stouter than neuro- 
podial bristles in Harmothoe) are distally entire and heavily serrated. 
Distribution. Previously recorded from Senegal and Angola. 

Family Chrysopetalidae 

Dorsal surface of the segments bearing a fan or transverse row of paleae. Pros- 
tomium with four eyes and three antennae. Biramous parapodia, with dorsal 
cirri on all segments. 

Genus BHAWANIA Schmarda, 1861 

The head is exceedingly small and not easily seen ; the elongated body consists of 
numerous segments. Neuropodial chaetae are compound. 

Bhawania goodei Webster, 1884 

Bhawania goodei, Augener, 1918, pp. 98-103, pi. 2, figs. 1-2 ; text-fig. 1 ; from Pram Pram. 

Localities, (i) Accra, Christiansborg shore (1), almost complete, 8 mm. long. 
(2) Axim, Hospital Reef, 7.i.5i (1), middle body piece, about 14 mm. long. 

It has not been possible to examine the head in either of these specimens, since, 
in the specimen from Accra, it has been damaged and it is missing from the Axim 
example. The neuropodial heterogomph spinigers and falcigers are as figured by 
Augener, but the paleae, which are of one type, are as shown in Text-fig. 3 and not 
as illustrated by Augener. Monro (1933) noted that the tips of the paleae were not 

Distribution. Apart from the above records B. goodei has been recorded from 
Bermuda and False Bay, South Africa, in the Atlantic Ocean, and from the Galapagos 
Islands and Taboga Island, Panama Bay, in the Pacific Ocean. 

Family Amphinomidae 

Body depressed, prostomium deeply set into the anterior segments, carrying three 
antennae, two palpal pads and a caruncle. Biramous parapodia with branchiae 
and simple chaetae (exceptionally uniramous with compound hooks). 

Key to Genera 

1. Body short and oval ; branchiae pinnate ....... Chloeia. 

Body long and vermiform ; branchiae bushy . . . . . . . 2. 

2. Branchiae beginning on the 1st chaetiger ; caruncle cushion-like, with dorsal surface 

with chevron crenulations ......... Hermodice. 

Branchiae beginning on the 2nd. or 3rd. chaetiger ; caruncle with a ventral crenulate 
portion Eurythoe. 

Genus CHLOEIA Savigny, 1818 
Caruncle a plaited crest on a horizontal plate. 



Fig. 3. Bhawania goodei. One of the paleae from the dorsal surface. 
(Specimen from Axim.) 

Chloeia viridis Schmarda, 1861 

Chloeia euglochis, Ehlers, 1887, pp. 18-24, pi- 1 > n S s - I_ ~ 2 > pi* 2 » n S s - I- 8 i pi- 3, n g s - I- 4- 
Chloeia euglochis, Augener, 191 8, pp. 94-95, from Goree, Senegal. 

Locality. Off Accra, Buchanan Survey, in 40 metres off Christiansborg (1), 
70 mm. long by 10 mm. wide. 

This is one of the few species of Polychaeta in which the pigment is retained, even 
after long periods in preservative fluid. Thus, in this single specimen, the dorsal 
cirri are violet, the pinnate gills, which appear at the fifth chae tiger, edged with 
purple, whilst the superior neuropodial chaetae are a brilliant orange, as are a few 
in the notopodial group. The remaining chaetae are glistening white, and the worm 
presents a striking appearance. The caruncle is attached to the body to the second 
chaetiger, though its unattached portion projects for 2-3 more segments, and it is 


surmounted throughout its length by a conspicuous, mid-dorsal, dark purple line. 
Another thin mid-dorsal purple line, which is slightly broken in the intersegmental 
furrows, extends along the back of the worm. 

Distribution. This species is also known from the W. Indies, Gulf of California, 
Galapagos and Cocos Islands. 

Genus HERMODICE Kinberg, 1857 
Caruncle cushion-like, with chevron crenulations on the dorsal surface. 

Hermodice carunculata (Pallas) var. didymobranchiata (Baird), 1864 

Amphinome didymobranchiata Baird, 1864, pp. 449-450, pi. 45, figs. 1-7 ; from Ascension 

Hermodice carunculata var. didymobranchiata, Fauvel, 1914, pp. 113-116, pi. 8, figs. 22-27, 

31-32 ; from the Gulf of Guinea. 
Hermodice carunculata var. didymobranchiata, Monro, 1930, p. 27 ; from the Gulf of Guinea 

and the French Congo. 

Localities. Off Accra : Stn. 47 (3), 12, 20 and 30 mm. long ; Stn. 115 (6), 
largest 40 mm. long, smallest 16 mm. ; Stn. 131 (1), 25 mm. long ; Stn. 133 (3), 
2 complete, about 20 mm. long. 

Most of these specimens have lost their colour except for the thin dark interseg- 
mental bands on the back. The caruncle extends to the 4th chaetiger. Separation 
of the branchiae into two bushy lobes instead of one and the almost complete absence 
of harpoon-shaped dorsal chaetae, separates this variety from the stem species. 

Distribution. This variety appears to have been recorded only from the above 
listed areas. 

Genus EURYTHOE Kinberg, 1857 

Caruncle with vertical folds along the lateral walls, and with a vertical crenulate 

Key to Species 

1. Branchiae beginning on the second segment. Caruncle terminating on the third 

or fourth segment. Four eyes ....... Ii. complanata. 

2. Branchiae beginning on the third segment. Caruncle terminating on the first 

segment. Four eyes ........ is. parvecarunculata. 

Eurythoe complanata (Pallas), 1766 

Eurythoe complanata, Augener, 191 8, pp. 88-89 ; from the Islands of Rolas and Annobon. 
Eurythoe complanata, Fauvel, 1932, pp. 45-46. 

Localities, (i) Tenpobo shore (70+). (2) Dixcove shore (7). (3) Axim, 
(a) Hospital Reef (1), posterior fragment ; (b) Shore (1), small specimen. 

These specimens vary considerably in length, the largest being about 100 mm. long, 
and the smallest 15 mm. 

Notopodial chaetae are of three types : (a) stout, straight and blunt ; (b) long and 
calcareous, with a slender tip having a slight spur at the base ; (c) long harpoon- 


shaped, with lateral rows of teeth, and neuropodial chaetae of two forms : (a) 
stout furcate chaetae with unequal arms ; (b) a few subfurcate with one arm thin and 

Distribution. This species is well known from littoral regions of the tropical 
Atlantic, Pacific and Indian Oceans. 

Eurythoe parvecarunculata Horst, 1912 

Eurythoe parvecarunculata, Augener 1918, pp. 90-93, pi. 2, fig. 3 ; pi. 3, figs. 37, 38, from 

Cameroons and Spanish Guinea. 
Eurythoe parvecarunculata, Fauvel 19276, pp. 525-526, fig. 1 a-h, from Cameroons and French 


Localities. Off Accra: Stn. 12 (1), anterior piece 17 mm. long; Stn. 16 (2), 
anterior pieces 10 mm. and 15 mm. long ; Stn. 32 (3), two 20 mm. long, one 30 mm. 
long ; Buchanan Survey, off Accra, in 3§-n metres, (1). 

There are stout, harpoon-shaped, and slender, elongated, bifurcated chaetae in 
the notopodial group. The bifurcation in the slender bristles may be reduced to a 
spur. The neuropodial chaetae are furcate, slightly denticulate. 

Distribution. This species is also known from the Indian Ocean, from Natal 
to Malaya. 

Family Phyllodocidae 

Body normally long and slender with numerous segments. Prostomium variable 
in outline, with two eyes and four or five antennae. Smooth or papillate proboscis, 
protrusible and unarmed. Segments 1-3 bearing tentacular cirri. Parapodia 
uniramous, with foliaceous dorsal cirri and compound chaetae. 

Key to Genera 

1. With 4 pairs of tentacular cirri .......... 2. 

With 2 pairs of tentacular cirri ......... Eteone. 

2. With 5 antennae ........... Eulalia. 1 

With 4 antennae ........... Phyllodoce. 

1 Not recorded from the Gold Coast. 

Genus PHYLLODOCE Savigny, 1822 

The four pairs of tentacular cirri are arranged on the three segments — 1 : 2 : 1. 
Two anal cirri. 

Key to Subgenera 

1. Protruded proboscis proximally set with longitudinal rows of papillae . . (Anaitides). 

2. Protruded proboscis proximally set with diffuse papillae . . . (Phyllodoce). 

Key to Species of Anaitides 

1. With more than 12 rows of papillae forming a continuous series around the proximal 

half of the proboscis .......... A. africana. 

2. With 12 rows of papillae arranged in two lateral groups of 6 on the proximal half of 

the proboscis . , , , , A . oculata. 


Phyllodoce (Anaitides) africana Augener, 1918 
Phyllodoce africana Augener, 1918, pp. 171-174, pi. 2, fig. 25 ; pi. 3, fig. 49-51 ; text-fig. 11. 

Locality. Off Accra, Stn. 52, (1), incomplete, 24 mm. long. 

The protruded proboscis is provided with a crown of six large bulbous papillae, 
from which run six longitudinal rows of eight similar papillae. These rows are 
followed by 24 longitudinal rows of about 14 much smaller papillae. Dorsal cirri 
broadly foliaceous throughout, ventral cirri slender, those situated posteriorly 
acquiring a pointed tip. 

Distribution. This species has been previously recorded only from Angola 
and Senegal. 

Phyllodoce (Anaitides) oculata Ehlers, 1887 

Phyllodoce {Anaitides) oculata, Augener, 1918, pp. 169-171, pi. 3, figs. 57-58, from Accra. 

This species has not been found in the present survey. 

Distally the proboscis is similar to the condition in A . africana, but the proximal 
portion has only 12 rows of papillae, and these are disposed in two lateral groups of 
six. Dorsal cirri become slender in the middle of the body. 

Distribution. The W. Indies, Gulf of Guinea, Ascension Island and Tristan da 

Genus ETEONE Savigny, 1822 

Dorsal cirrus absent on the 2nd segment. Proboscis smooth, or with soft papillae 
and small cuticularized tubercles. 

Key to Subgenera 

1. Proboscis smooth or with soft papillae ....... (Eteone). 

2. Proboscis with lateral rows of large soft papillae and small spinous tubercles . (Mysta). 

Eteone (Mysta) picta Quatrefages, 1865 

Eteone {Mysta) picta, Fauvel, 1923, pp. 176-177, fig. 64, a-g. 

Localities. Off Accra : Stn. 57 (1), 15 mm. long, incomplete ; Stn. 69 (2), 1 
complete 17 mm., the other incomplete, 37 mm. long ; Buchanan Survey, 3-6 to 
ii-o metres (4), 3 complete, 31, 38 and 42 mm. long. 

For description these specimens may conveniently be divided into two groups : 
one representing the young or immature form, the other the adult or sexually mature 

(a) The immature forms (from Stns. 57 and 69). In these specimens the prostom- 
ium bears two clear indentations on either side. The protruded proboscis is crowned 
by an encircling group of large soft papillae, whilst two longitudinal rows of equally 
large papillae extend laterally along the greater part of its length. Elsewhere there 
are small papillae with soft chitinous teeth (Text-fig. 4, a). The dorsal surface of 
the body in the specimen from Stn. 69 is marked with pale red-brown lateral stripes, 



although the effect of preservation may give the uniform stripes a speckled appear- 
ance. In the specimen from Stn. 57 the colour is uniformly light brown, with dark 
markings towards the sides. All specimens have the base of the foliaceous dorsal 
cirrus coloured in pattern with the colouration of the adjacent dorsal surface of the 
body. The remainder of the dorsal cirrus is greyish white or yellow, there being a 
suture between these two clearly defined areas. Compound chaetae have long ter- 
minal articles and project much further from the body than the cirri. On the articu- 

Fig. 4. Mysta picta. (a) Small papilla, with chitinous teeth, from the proboscis, x 700 
(b) compound chaeta, x 500. (Specimen from Stn. 69.) 

lating surface of the main stem of the chaetae there are two teeth, as well as a small 
finely pectinate plate (Text-fig. 4, b). 

(b) The sexually mature forms (Buchanan Survey collections). In two of these 
specimens the colour pattern is similar to that in the specimens described in group 
(a), but the posterior parapodia contain small eggs, in which the nuclei are not visible. 
The suture on the dorsal cirrus, however, is clearly seen and appears to segregate 
a basal sac-like portion from a terminal solid piece. One of the remaining specimens 
has much longer dorsal cirri extending posteriorly from the middle of the body. 
In the anterior and middle regions of the dorsal surface the body is without colour 


but gradually becomes light bronze posteriorly. The lateral body-wall and the basal 
sac-like portion of the parapodia, from behind the middle region of the body are rich 
blood red, whilst the tips of the dorsal cirri are yellow. The basal portion contains 
numerous large eggs, with visible nuclei ; the eggs are so large and numerous that 
they produce a considerable swelling of the body- wall. The remaining specimen 
has very much enlarged dorsal cirri from the 37th chaetiger, at which point the dorsal 
body-wall and the bases of the parapodia have assumed a bronze colour, whilst the 
tips of the dorsal cirri are yellow. Anterior to the 37th chaetiger the colour is similar 
to that in specimens described in group (a). The parapodia, posteriorly from the 
37th chaetiger, all contain eggs, the number to each segment increasing towards the 
hinder end of the body, whilst the chaetae only project as far as the dorsal cirri. 
Constant reference has been made to the prominent suture on the dorsal cirrus in 
all the specimens here described, and examination of the mature specimens indicates 
that the eggs collect in the basal sac-like portion of the parapodia, and it is here 
suggested that the suture may represent the line of rupture when the eggs are liber- 
ated through the body wall, into the sea, for external fertilization. 

That these specimens represent a series from the immature to the sexually mature 
condition is a supposition which can only be confirmed through observation on the 
living animal. But the conclusion here reached — that all the specimens are of the 
same species — appears justifiable, particularly when the variation in related genera 
and families during their life histories is considered. 

Distribution. This appears to be the first record of the species from the Equa- 
torial Atlantic. Previously, E. (M.) picta has been reported from the North Sea, 
the English Channel, the west coast of Scotland, the Bay of Biscay and the Mediter- 

Family Hesionidae 

Body short, cylindrical. Prostomium simple or bilobed, normally with four eyes, 
two or three antennae and two biarticulate palps. Protrusible proboscis with or 
without jaws. Parapodia biramous, or reduced with long dorsal cirri and both 
simple and compound chaetae. 

Genus LEOC RATES Kinberg, 1865 

Prostomium with four eyes, three antennae and two palps. There are eight pairs 
of tentacular cirri. Proboscis with chitinous jaws. Parapodia biramous, noto- 
podial chaetae simple, neuropodial chaetae compound. 

Leocrates claparedii (Costa), 1868 

Tyrrhena claparedii, Claparede, 1868, pp. 228-231, pi. 18, fig. 3. 
Leocrates claparedii, Fauvel, 1923, pp. 237-238, fig. 88, i-n. 

Locality. Axim, Hospital Reef, 13. iv. 49 (1), complete, 18 mm. long. 

In this species the upper jaw plates have only one tooth, which distinguishes it 
from L. atlanticus, with two teeth, reported by Fauvel (1950) from Dakar. The 
median antenna in L. claparedii is short and subulate ; the paired laterals slender 


and slightly longer than the palps. Notopodial chaetae appear at the 5th chaetiger 
and are simple and spinous : neuropodial chaetae have a bidentate sickle-shaped 
terminal piece. There are 17 segments bearing parapodia, the last pair being 
achaetous, retaining only the notopodial and neuropodial cirri as parapodial append- 
ages, which, with the two urites, gives the appearance of there being six anal cirri. 

Augener (1918) reported a new species, L. greeffianus, from the Isle of Rolas, by 
St. Thomas, without noting the form of the pharyngeal armature. 

Distribution. Originally recorded from Naples ; Day (1934 and 195 1) has 
collected this species from the coast of Natal. It may have been found in the 
Indo-Pacific region and reported under various names, but lack of details of the jaw 
structure makes confirmation of this impossible. 

Family Syllidae 

Body small and narrow. Prostomium with four eyes, two palps and three anten- 
nae. Normally two pairs of tentacular cirri on the first segment, which never has 
chaetae. Protrusible proboscis, divided into two regions : (1) anterior, the pharynx, 
chitinous, cylindrical, with or without teeth ; (2) posterior, the proventriculus, barrel- 
shaped. Parapodia uniramous, normally with dorsal and ventral cirri. Chaetae 
rarely simple, normally compound with a heterogomph articulation. 

Key to Subfamilies 

1. Without ventral cirri ......... Autolytinae. 1 

With ventral cirri ............ 2. 

2. Palps not fused ........... Syllinae. 

Palps fused ............. 3. 

3. Palps fused at the bases only ........ Eusyllinae. 

Palps fused along their entire length ....... Exogoninae. 

1 Not recorded from the Gold Coast. 

Subfamily Syllinae 
Key to Genera 

1. A large tooth with a trepan ........ Trypanosyllis. 

A large tooth with no trepan .......... 2. 

2. With the tooth on the anterior part of the pharynx ..... Syllis. 
With the tooth on the posterior part of the pharynx .... Opisthosyllis. 

Genus TRYPANOSYLLIS Claparede, 1864 

The proboscis is crowned with a circle of chitinous teeth (the trepan), and there is 
also a large single anterior tooth. 

Trypanosyllis prampramensis Augener, 1918 

Trypanosyllis prampramensis Augener, 1918, pp. 276-278, pi. 4, figs. 91-92, text-fig. 26 ; from 

Pram Pram. 
Trypanosyllis prampramensis, Day, 1953, p. 414, from Still Bay and Kommetje, S. Africa. 

This species is only known from these records, and has not been found in the present 
survey. T. prampramensis appears to be very close to the European T. coeliaca, 


but is characterized by unidentate compound chaetae, the latter having bidentate 
chaetae. Augener's specimen was 6-5 mm. long by 0-7 mm. wide for 100 segments. 

Genus SYLLIS Savigny, 18 18 

Prostomium with four eyes and often two anterior eye-spots. Pharynx crowned 
with soft papillae and with a prominent anterior tooth. Antennae and dorsal cirri 
are moniliform ; ventral cirri are not articulated. 

Key to Subgenera 

1. All chaetae simple .......... (Haplosyllis). 1 

With compound chaetae and, sometimes, simple chaetae . . . . . 2. 

2. Anterior chaetae compound, thereafter some large simple chaetae . . . (Syllis). 
All chaetae compound . . . . . . . . . . . 3. 

3. Compound chaetae with short articles all alike or differing only slightly from each 

other ............ (Typosyllis). 

Compound chaetae with long and short articles . . . . . (Ehlersia) . 

1 Not recorded from the Gold Coast. 

Syllis (Syllis) gracilis Grube, 1840 

Syllis gracilis, Fauvel, 1923, p. 259, fig. 96, f-i. 

Locality. Off Accra, dredge in 7-3 metres, 1 mile off shore, 2 miles west of 
Densu River (1), 8 mm. long. 

The dorsal cirri have between 8 and 12 articles. In the middle of the body chaetae 
are simple (Text-fig. 5, a), but anteriorly and posteriorly there are compound falcigers. 
The specimens here examined are not as long as those recorded by Fauvel (1923) ; 
they may be immature. 

Distribution. Mediterranean Sea, Atlantic Ocean, Red Sea, Indian and Pacific 

Subgenus TYPOSYLLIS Langerhans, 1879 
Key to Species 

1. Dorsal cirri long with more than 20 articles ..... T. variegata. 

2. Dorsal cirri short with less than 20 articles . . . . . . . T. hyalina. 

Syllis (Typosyllis) variegata Grube, i860 
Syllis {Typosyllis) variegata, Fauvel, 1923, p. 262, fig. 97, h-n. 

Localities. Axim, (a) Shore, 13.iv.49 (1), 10 mm. long, incomplete; (b) 
Hospital Reef, 7.i.5i (1), 7 mm. long, complete. 

In the specimen from Axim shore, the dorsal cirri are alternately long and short, 
with 30 to 40 and 20 to 25 articles respectively. The specimen from the Hospital 
Reef, however, has dorsal cirri all about the same size with 30 to 40 articles. In 
both specimens the pharynx is crowned with 10 to 11 soft papillae and has a promi- 
nent anterior tooth. In the specimen from the Hospital Reef compound chaetae 


have strongly bidentate, short, terminal articles, but in the specimen from the shore 
the bidentation is less pronounced. 

Distribution. English Channel, Mediterranean Sea, Atlantic Ocean, Indian 
Ocean, Red Sea and Pacific Ocean. 

Syllis (Typosyllis) hyalina Grube, 1863 

Syllis (Typosyllis) hyalina, Augener, 1918, pp. 242-247, pi. 4, figs. 95, 96, from Pram Pram. 

This species has not been collected in the present survey. It is distinguished 
from T. variegata in having unidentate terminal articles on the compound chaetea, 
and few articles on the dorsal cirri. T. hyalina frequently retains its colour after 
preservation, showing a uniform light red anteriorly, or transverse bands of a red- 
brown, but no specimens of T. variegata have been examined retaining any colouring 

Distribution. English Channel, Mediterranean Sea and Atlantic Ocean. 

Subgenus Ehlersia Langerhans, 1879 

Syllis (Ehlersia) cornuta Rathke, 1843 

Syllis (Ehlersia) sexoculata, Augener, 191 8, pp. 269-271, from Pram Pram in 9 metres depth. 

Localities, (i) Off Accra, dredge in 7-3 metres, 1 mile off shore, 2 miles west 
of Densu River, (1) , 6 mm. long. (2) Axim shore, 13 /14 . iv . 49, (2) , 5 and 6 mm. long. 

There are two pairs of small eyes posterior to the median antennae, and two 
minute eye-spots on the anterior border of the prostomium. Anterior dorsal cirri 
have 8 to 10 articles, median cirri 10 and posterior cirri 4 to 5. The two anal cirri 
have 12 articles. Compound chaetae are of two kinds, one with a long terminal 
article (Text-fig. 5, b) and the other short (Text-fig. 5, c). 

These specimens are small and may be juveniles. 

Distribution. English Channel, Mediterranean Sea, Atlantic — Canaries, Senegal, 
Angola — Persian Gulf and Indian Ocean. 

Genus OPISTHOSYLLIS Langerhans, 1879 
An occipital flap on the posterior border of the prostomium. 

Opisthosyllis brunnea Langerhans, 1879 
Opisthosyllis brunnea, Augener, 1918, pp. 274-276, text-fig. 25. 

Localities, (i) Tenpobo shore, 3/4.U.50 (5), 3 complete, 8-10 mm. long. 
(2) Winneba shore, 22.xi.49 (1), complete, 13 mm. long. (3) Axim, Hospital 
Reef, 7.L51 (1), incomplete, 6 mm. long. 

There are between 30 to 50 articles in the dorsal cirri. The pharynx extends over 
eleven segments, bears a crown of 9 to n papillae and carries a large conical tooth 





Fig. 5. Syllidae chaetae. (a) Syllis gracilis, simple chaeta, x 300. (b) Ehlersia 
cornuta, compound spiniger, x 400. (c) Ehlersia cornuta, compound falciger, x 400. 
(d) Opisthosyllis brunnea, compound falciger, x 250. (e) Opisthosyllis brunnea, 
simple chaetae, x 300. 

posteriorly. The tooth is without colour and stands out against the dark brown of the 
pharynx. Compound chaetae have short unidentate articles (Text-fig. 5, d). In 
posterior parapodia there are long simple chaetae (Text-fig. 5, e). 

Distribution. The tropical coast of W. Africa and the Indian Ocean. 

Subfamily Eusyllinae 
Genus PIONOSYLLIS Malmgren, 1867 
Pharynx with a single anterior tooth. 

ZOOL. 3, 2. 


Pionosyllis ehlersiaeformis Augener, 1913 

Pionosyllis ehlersiaeformis, Augener, 1918, pp. 281-283. 
Pionosyllis ehlersiaeformis, Day, 1953, pp. 415-416, text-fig. 3, d. 

Locality. Off Accra, Buchanan Survey, in 14-5 metres off the Castle (1), in- 
complete, 15 mm. long. 

The pharynx is crowned with eight large papillae. Antennae and dorsal cirri 
are very long with numerous articulations, which is unusual in this genus where, 
normally, the appendages are smooth. Ventral cirri are smooth and pointed. 
Superior chaetae have slender, curved unidentate terminal articles, about ten times 
as long as the articles on the inferior chaetae which are straight with bidentate tips. 

No eyes have been seen in this specimen, otherwise it is typical of the species. 

Distribution. Augener originally recorded this species from S.W. Australia, but 
in 1918 he reported it from German S.W. Africa, and in 1953 Day collected it from 
Cape Agulhas. 

Subfamily Exogoninae 

Genus EXOGONE Oersted, 1845 

Palps well developed, completely fused ; three antennae ; one pair of tentacular 
cirri. Pharynx with a single tooth. Dorsal and ventral cirri small. Chaetae 
simple and compound. 

Exogone getnmifera (Pagenstecher) 

Exogone gemmifera, Augener, 19 18, pp. 299-301, text-fig. 29, from Pram Pram in 9 metres 

This species has not been collected in the present survey. Median antenna 
longer than the prostomium and longer than the paired laterals ; two pairs of large 
eyes. Dorsal cirri missing on the second chaetiger ; ventral cirri small, often difficult 
to see. Pharynx straight, crowned with papillae, with a single anterior tooth. 
In each parapodium there is a simple, bluntly terminated chaeta and a compound 
form with a very small bidentate article ; on posterior feet there is a simple ventral 
chaeta. E. gemmifera normally measures between 2-4 mm. in length. 

Distribution. English Channel, Mediterranean Sea, and Atlantic Ocean. 

Family Nereidae 

Prostomium with four eyes, two antennae, two massive palps and four pairs of 
tentacular cirri. Proboscis armed with two horny jaws and normally a series of 
horny teeth (paragnaths). Parapodia normally biramous after the 2nd foot ; 
chaetae compound ; most species have a Heteronereis, sexually mature form. 

Generic and specific distinctions in this family are based primarily on the form 
and arrangement of the paragnaths. These are divided first into two rings, the 
maxillary and oral (Text-fig. 6), and then into eight numbered areas, of which I to IV 



are maxillary and V to VIII oral ; odd numbers are median and single, even numbers 
lateral and paired. The jaws are at the distal end of the proboscis. The different 
types of chaetae referred to in the text are illustrated in Text-figs. 8 and 12. 




(a) W 

Fig. 6. Diagrammatic representation of the division of the proboscis in Nereidae. 

Key to Genera 

1. Horny paragnaths of one type only, conical ..... 
Horny paragnaths of more than one type ...... 

2. Horny paragnaths of two types, conical teeth and transverse cutting plates 
Horny paragnaths of three types, conical, transverse and pectinate 



. Perinereis. 


Genus NEREIS Linnaeus, 1758 

All groups of paragnaths complete — Neanthes, sub-genus. 

One or more groups of paragnaths missing — Nereis, sensu stricta. 

Nereis falsa Quatrefages, 1865 

Nereis callaona, Augener, 19 18, pp. 184-186. 

Nereis falsa, Fauvel, 1923, pp. 337-338, rig. 129, e-m. 

Nereis falsa, Day, 1951, pp. 27-28. 

Localities, (i) Tenpobo shore (3), anterior pieces only, n-15 mm. long. (2) 
(a) Accra, Christiansborg shore, 17. hi. 49 (20 -f), many complete, between 30-60 
mm. long ; (b) Accra, on sponge near Sound, 9.xii.5o (1), anterior fragment 10 mm. 
long ; (c) from wood netted off Chorkor, 14. v. 51 (1), complete, 18 mm. long. (3) 
(a) Winneba shore (1), anterior fragment, 10 mm. long ; (b) Winneba, neap tide, 
15.xi.49 (3), 20-30 mm. long. (4) Axim, (a) Shore 13/14.^.49 (4), 2 complete, 
50 mm. long, others anterior fragments, 10-15 mm - l° n g '> (&) Hospital Reef, 7.L51 



(9), some complete, 10-15 mm - l° n g > i c ) Lighthouse Reef, 8.L51 (2), complete 10 
and 20 mm. long. 

This is one of the commonest littoral forms of the area. The arrangement of the 
paragnaths varies very little from that shown in Text-fig. 7, though paired groups 
VI sometimes have three or five teeth instead of two sets of four ; area V never has 
any paragnaths. Notopodial chaetae are homogomph spinigers throughout, except 



Fig. 7. Nereis falsa. Arrangement of the paragnaths on the proboscis, 
(a) dorsal ; (b) ventral. (Specimen from Tenpobo.) 

that in some of the posterior feet a single homogomph falciger may be present 
(Text-fig. 8, a). Neuropodia have homogomph spinigers and heterogomph falcigers 
dorsally and heterogomph spinigers (Text-fig. 8, b) and falcigers ventrally. 

Distribution. — Atlantic, Mediterranean, Adriatic and around the South African 
coast from Cape Town to Natal. 

Nereis (Neanthes) succinea (Frey & Leuckart), 1847 

Nereis glandulosa, Augener, 1918, pp. 192-194. 

Nereis succinea Fauvel, 1936, pp. 307-313, {nee Nereis succinea Fauvel, 1923, pp. 346-347.) 

Localities. Off Accra : (a) From wood netted of Chorkor, in 14 metres, 14. v. 51 
(3), complete, 11-25 mm - l° n g '> (&) Stn. 29 (1), complete, 25 mm. long ; (c) 12 metres 
off Accra (1), 38 mm. almost complete ; (d) Buchanan Survey, in a depth of 3-6 to 
ii- o metres (10), between 25-60 mm. long, one a heteronereis. 

The pattern of paragnaths varies considerably in this species around a typical 



form shown in Text-fig. 9. Thus I =1-4 in a longitudinal line ; II = arcs in two 
rather scattered rows ; III = a transverse group in 2-3 scattered rows ; IV = arced 
masses ; V = 0-4 (rarely 5-7). The absence of teeth on V in four of the specimens 
from the Buchanan Survey does not necessarily upset the validity of the above key, 
since this may be due to immaturity, accidental loss, general wear and tear, or it 
may indicate the approach of sexual maturity. This latter reason is of doubtful 
significance, but is referred to because this group is missing in the heteronereis 

Fig. 8. Nereis falsa, (a) Homogomph falciger from a posterior notopodium, (b) hetero- 
gomph spiniger from a neuropodium. Both X 250. (Specimen from Axim.) 

example. Area VI is typically described as having a circle of six to seven paragnaths 
surrounding a central one, but the majority of specimens here examined conform 
to the pattern shown in Text-fig. 9, in which between 9-13 teeth are arranged in 
three scattered rows. A specimen in the B.M. (N.H.), Reg. No. 1928.4.26.290, 
from French Guinea, identified by Professor Fauvel, has 9-10 teeth in three scattered 
rows, and this variation may be common among W. African examples of N. succinea. 
Teeth in areas VII and VIII are both small and large and arranged, approximately, 
in two rows. 

A parapodium from the posterior region of one specimen is shown in Text-fig. 10. 
The enlargement of the dorsal lobe only takes place posteriorly, anteriorly it is 
long and thin. Notopodial chaetae are homogomph spinigers, neuropodial chaetae 



Fig. g. Nereis succinea. Arrangement of the paragnaths on the proboscis, (a) 
dorsal, (b) ventral. (Specimen from the Buchanan Survey.) 



homogomph spinigers, and heterogomph spinigers and falcigers. The absence of 
notopodial homogomph falcigers is considered by many workers (Fauvel, 1936 ; 
Hartman, 195 1) to be important diagnostically. It is doubtful if a negative charac- 
ter of this type can be so considered, since accidental loss of chaetae frequently occurs. 
Distribution. Cosmopolitan ; frequently found in waters of low salinity. 

0.5 mm 

Fig. 10. 

Nereis succinea. Posterior view of parapodium from the 32 nd chaetiger. 
(Specimen from the Buchanan Survey.) 

Genus PERINEREIS Kinberg, 1865 

With transverse chitinous plates (horny paragnaths) on area VI ; other areas with 
conical paragnaths. 

Perinereis melanocephala Mcintosh, 1885 

Nereis (Perinereis) melanocephala, Mcintosh, 1885, pp. 216-219, pi. 34 figs. 14-17 ; pi. i6a, 

figs. 8, 9 ; from Bermuda, between tide marks. 
Nereis (Perinereis) melanocephala, Augener, 1918, pp. 209-212, from the Island of Annobon and 

the Cameroons. 

Localities, (i) Tenpobo shore (8), 6 complete, 20-30 mm. long, others ant. 
fragments : collected February, 1950. (2) Accra, Christiansborg shore (1), ant. 
piece 38 mm. long. (3) Winneba shore, 15.xi.49, Neap tides (2), anterior pieces, 
17-20 mm. long — one a developing heteroneis stage. (4) Sekondi, high level plateau, 
21.ii.49 (3) complete, 20-25 mm. long. (5) Dixcove shore (1), complete, 64 mm. 
long. (6) Axim, 13/14.1V.49 (1), complete, 18 mm. long. 

9 8 


The proboscis of this species is shown in Text-fig. II. Little variation from the 
normal has been found ; I = a triangular area of teeth ; II = rhomboidal area of 
similar size to I ; III = rectangular area also equal to I ; IV similar to II ; V = 
a single large tooth ; VI = single long, transverse, horny bands ; VII and VIII = 
2 rows of teeth with scattered isolated teeth here and there. This compares accur- 
ately with Mcintosh's original description. Unfortunately the type specimen, 
B.M. (N.H.). Reg. No. 1885. 12. 1. 159 has no head. Efforts to find this portion, 
including enquiries at the University of St. Andrews, where Mcintosh did his work, 
have been unsuccessful. Notopodial chaetae are all homogomph spinigers (Text-fig. 


Fig. 11. Perinereis melanocephala. Arrangement of the paragnaths on the proboscis, 
(a) dorsal, (b) ventral. (Specimen from Sekondi.) 

12, b), as are the superior neuropodial group. Inferior neuropodial chaetae are hetero- 
gomph falcigers (Text-fig. 12, a). The superior notopodial lobe becomes enlarged 
in the region of the 37th foot, continuing thus to the end of the body, the dorsal 
cirrus being attached (Text-fig. 13, a) to its indented extremity. The degree of this 
enlargement may be noted with respect to an anterior foot illustrated in Text-fig. 


One of the specimens from Winneba has modified parapodia, beginning at the 17th 
segment, indicating the development of the heteronereis stage (Text-fig. 14). The 
lobe at the base of the dorsal cirrus, the post-chaetal neuropodial lobe and the ventral 
cirrus are all in the early stages of development into foliaceous swimming surfaces. 
The chaetae, however, remain normal, not having developed into the natatory type 
expected in this genus, otherwise the aspect of the parapodia is similar to that 
assumed by the more common P. cultrifera in the heteronereis condition. 

Distribution, P, melanocephala is only known from the above records. 



Fig. 12. Perinereis melanocephala. (a) Neuropodial heterogomph falciger, x 500 ; (b) 
notopodial homogomph spiniger, x 250. (Specimen from Sekondi.) 

Genus PSEUDONEREIS Kinberg, 1865 

With transverse chitinous plates (horny paragnaths) on area VI ; transverse rows 
of pectinate teeth and conical paragnaths on other areas. 

Pseudonereis gallapagensis Kinberg, 1865 

Pseudonereis gallapagensis, Gravier, 1909, pp. 629-633, figs. 
Pseudonereis variegata, Fauvel, 1927b, pp. 527-528, (part). 
Pseudonereis gallapagensis, Hartman, 1948, pp. 68-69. 

Localities, (i) (a) Winneba shore, 3 . iii . 50 (1), complete, 9 mm. long, 4.^.50(2), 
one complete, 15 mm.; (b) Winneba rock face, 22.xi.49 (6), several complete, 



Fig. 13. Perinereis melanocephala. Parapodia of (a) 37th chaetiger (specimen from 
Sekondi) ; (b) an anterior chaetiger (specimen from Tenpobo). 



Fig. 14. Perinereis melanocephala. Parapodium from the 23rd chaetiger of a developing 
heteronereid. (Specimen from Winneba.) 


between 15-30 mm. long; (2) (a) Axim shore, 13/14.1V.49 (1) complete, 19 mm. 
long ; (b) Axim, hospital reef (2), both complete, one approaching sexual maturity, 
17 mm. long, the other n mm. long. 

Notopodial chaetae are homogomph spinigers; superior neuropodial chaetae 
also homogomph spinigers, whilst the inferior group are heterogomph falcigers. 
Posteriorly the dorsal lobe becomes foliaceous and carries at its distal end the dorsal 
cirrus, leaving little or no free margin. Hartman (1948) referred to this character 
as the specific distinction between P. gallapagensis and P. variegata (Grube), noting 
that the latter species had a free portion at the tip of the dorsal foliaceous lobe. 
Day (195 1) disputed the value of this distinction and it is doubtful if it is of practical 
use. It is here suggested that P. gallapagensis has on each area IV of the proboscis 
an apical group of about 16 small conical teeth in addition to the rows of pectinae 
(Text-fig. 15, c), and this is a diagnostic character separating it from P. variegata. 
Kinberg (1865) did not note this character, though his figures were not drawn in a 
position which would show it. In examining the type specimen, which was in poor 
condition, Hartman (1948) did not refer to the proboscidean structures. Gravier 
(1909), however, observed this apical group on IV in specimens collected from Peru. 

In groups VII and VIII in P. gallapagensis there is one row of alternating long and 
short teeth (Text-fig. 15, b), the anterior tip of the long teeth being in line with the 

Fig. 15. Pseudonereis gallapagensis. Arrangement of the paragnaths on the proboscis, 
(a) dorsal, (b) ventral, (c) apical (jaws left out). (Specimen from Axim.) 

latter. This is the condition on an extruded proboscis. If not extruded, however, 
due to the different lengths of alternate teeth, and the general contracted condition 
of the proboscis, it appears as if two rows of teeth are present. This character is 
clearly shown in Kinberg's (1910) type figure and contrasts with the condition in 
P, variegata in which there are 2/3 rows of short teeth (Mcintosh, 1903). All speci- 


mens here listed agree with this description, as does one from the Cameroons in the 
B.M. (N.H.), Reg. No. 1928.4.2.844, which was recorded by Fauvel, 19276, as 
P. variegata. Further specimens of P. gallapagensis in the B.M. (N.H.) collections 
come from Ceylon and from Mormugas Bay, Goa, near Bombay. 

Distribution. This is the first record of this species from the W. African coast ; 
apart from the above records, P. gallapagensis is known widely from the western 
coast of S. America, the Galapagos Islands and Hawaii. 

Family Nephthydidae 

Body with numerous short segments. Prostomium small and flat carrying two 
pairs of antennae. Proboscis terminated with 14 to 22 soft bifid papillae and bearing 
14 to 22 longitudinal rows of similar processes externally and two horny jaw pieces 
on its inner wall. Parapodia are biramous except for the first pair which may not 
be fully developed ; rami are wide apart with a coiled cirrus, probably branchiate 
in function, between them. 

Only three genera are recognized in this family and the following key has been 
adapted from Hartman (1950) in which work an acceptable division of the type 
genus Nephthys was suggested : 

1. Inter-ramal cirri (branchiae) recurved — evolute ...... Nephthys. 

2. ,, „ ,, involute ....... Aglaophamus. 

3. ,, ,, ,, absent ....... Micronephthys. 

No examples of the genus Micronephthys Freidrich, 1939, have been collected 
from the Gold Coast, but Augener (1918) recorded M. ambrizettana from Angola. 

Genus NEPHTHYS Cuvier, 1817 
Nephthys hombergii Audouin & Milne-Edwards, 1830 
Nephthys hombergii, Fauvel, 1923, p. 367, fig. 143, a-d. 

Localities. Off Accra : Stn. 71 (1) 12 mm. long, anterior piece ; Stn. 132 (1) 
16 mm. long, anterior piece ; Buchanan Survey, in 3-5 to 11 metres (2), one complete, 
54 mm. long. 

This well-known species is characterized by 22 longitudinal rows of papillae on 
the proboscis, the appearance of the branchiae on the 4th chaetiger and the presence 
of a notopodial button ventral to the acicula and tending to overgrow it. 

Distribution. North Sea, English Channel, Mediterranean, Atlantic. 

Genus AGLAOPHAMUS Kinberg, 1865 
Aglaophamus lyrochaetus (Fauvel), 1902 

Nephthys lyrochaeta Fauvel, 1902, pp. 72-5, figs. 9-12, from the estuary of the R. Casamance, 

Nephthys lyrochaeta, Augener, 1918, pp. 160-166, pi. 2, fig. 12 ; pi. 3, fig. 59, from Saltpond, 

Gold Coast in 9 metres, and Accra ; also several other W. African stations, 


Localities. Off Accra: Stn. 32 (1), anterior piece, 10 mm. long; Stn. 59 (1), 
anterior piece, 7 mm. long ; Stn. 70 (2), both anterior pieces, 7 and 8 mm. long ; 
Buchanan Survey, in 3-5-11 metres depth (4), two complete, 26 and 28 mm. long. 

There are 14 longitudinal rows of papillae on the proboscis in this species. The 
" lyre " bristles occur in the centre of each group of chaetae and are about half the 
length of the others. They are therefore difficult to see unless the parapodium 
chosen for observation is carefully mounted. A later diagnosis by Fauvel (19276) of 
A . lyrochaetus indicated the presence of a small lobe on the dorsum of the neuropo- 
dium which was confirmed by Monro (1930). With the " lyre " bristles, this character 
serves as specifically diagnostic. The anterior feet are biramous and not uniramous 
as indicated by Augener (1918), and the long ventral and dorsal cirri of the first 
and second chaetigers respectively, noted by Monro (1930), are not present in the 
specimens here described. 

Distribution. — A . lyrochaetus is only known through the above records. 

Family Glyceridae 

Body elongate, tapering to both extremities, segments bi- or triannulate. Prosto- 
mium conical, annulated, with four small terminal antennae. Protrusible proboscis, 
very long, covered with papillae and armed with horny jaws of various kinds. 
Normally with biramous parapodia, but frequently uniramous anteriorly. Branchiae, 
when present, compound or simple, often retractile. Chaetae simple or compound. 

Key to Subfamilies 

1. Body divided into 2 or 3 regions by variation in the form of the parapodia which 

may be uniramous, sub-biramous or biramous. Jaws and paragnaths numerous 


2. Body not divided into distinct regions, parapodia either uniramous or biramous 

throughout. Jaws and paragnaths few in number .... Glycerinae. 

Subfamily Glycerinae 
Key to Genera 

1. Parapodia uniramous throughout, with compound cheatae only . . Hemipodus. 1 

2. Parapodia biramous throughout, with simple chaetae in notopodia and compound 

chaetae in neuropodia .......... Glycera. 

1 Not recorded from the Gold Coast. 

Genus GLYCERA Savigny, 1818 

Prostomium long, with more than three annulations. Proboscis with four large 
horny paragnaths distally and covered with papillae. Parapodia with a stumpy 
dorsal cirrus, two anterior lobes one or two posterior lobes and a ventral cirrus. 
Neuropodial chaetae compound spinigers, notopodial chaetae simple capillaries. 
Branchiae present or absent, simple or branched, permanent or retractile into the 



Glycera convoluta Keferstein, 1862 

Glycera africana, Arwidsson, 1898, pp. 21-22, pi. 1, figs. 10-12. 

Glycera africana, Augener, 1918, pp. 384-386. 

Glycera tridactyla, Augener, 1918, pp. 386-389, pi. 5, figs. 142-143, text-fig. 47. 

Glycera convoluta, Fauvel, 1923, pp. 383-385, fig. 150, a-h. 

Localities, (i) Off Accra: Stn. 65 (1); Stn. 69 (1) ; Stn. 89 (1) ; Stn. 126 (1) ; 
all incomplete except Stn. 65, which measures 55 mm. long ; Buchanan Survey in 
3- 6 to ii- o metres depth (6), five complete, 40-52 mm. in length. (2) Apam shore, 
3/4.^.50 (3), one complete, 46 mm. long. 

The papillae of this species are illustrated in Text-fig. 16, a and b. They are 
cylindrical with a truncated termination, the truncation bearing a plate similar in 


Fig. 16. Glycera convoluta. (a) Papilla from the proboscis, en face view, x 1500 (speci- 
men from Apam) ; (b) papilla from the proboscis, side view, x 1500 (specimen from 
Stn. 89) ; (c) silhouette of one of the 4 jaw articles, drawn in situ, (specimen from 
Stn. 126). 

appearance to the nail on the human finger. Over the greater part of the body both 
anterior parapodial lobes are bluntly pointed and each project the same distance 
from the body ; posterior lobes are unequal, the dorsal being blunt to oval and 
projecting a short distance, whilst the ventral lobe is blunt and hardly projects at 
all. Branchiae are non-retractile and lie on the dorsal edge of the parapodia. In 
the anterior region they project as far as the chaetae, but posteriorly become very 
much longer. Normally the branchiae appear at the I4th-i8th parapodia, but they 
may not appear until the 29th foot. They are absent from extreme posterior segments. 

A silhouette of one of the paragnaths is shown in Text-fig. 16, c, it was drawn in 
situ, at the end of the protruded proboscis. 

Distribution. Mediterranean Sea and Atlantic Ocean. 


Subfamily Goniadinae 

Key to Genera 

1. Proboscis with chevrons .......... Goniada. 

Proboscis without chevrons . . . . . . . . . . 2 . 

2. Neuropodia with spinigerous chaetae only ...... Ophioglycera, 

Neuropodia with spinigerous and falcigerous chaetae .... Goniadopsis, 

Genus GONIADA Audouin & Milne-Edwards, 1833 

Body divided normally into two regions, an anterior uniramous and a posterior 
biramous portion ; there may be a transitional region with sub-biramous parapodia. 
Paragnaths include a pair of large dentate macrognaths separated from each other 
by dorsal and ventral arcs of micrognaths. The proboscis carries two rows of dark 
hard V shaped pieces, chevrons, and numerous papillae of one or few kinds, dis- 
tributed in longitudinal, or irregular bands. 

Goniada multidentata Arwidsson, 1899 

Goniada multidentata Arwidsson, 1898, pp. 45-47, figs. 40-42. 

Locality. Off Accra, Buchanan Survey in 39-6 metres depth, off Lagoon (1), 
complete, about 90 mm. long. 

There are 36 anterior uniramous, 15 median sub-biramous, and 131 posterior 
biramous, parapodia. The uniramous feet bear prominent thick ventral cirri and 
slender pointed dorsal cirri. There are three ligules to each chaetigerous lobe, two 
anterior, long and finger-shaped, and one posterior, short. The sub-biramous portion 
is so qualified only because the notopodial lobes are not well developed and also 
because the segments in this region are the same width as those in the anterior 
uniramous region. Notopodial chaetae, however, are developed and the segments 
are strictly biramous. In the biramous posterior portion the segments are much 
wider than anteriorly. Neuropodial ligules are as in the anterior region except that 
the posterior ligule is triangular ; there are two notopodial ligules, both blunt, one 
dorsal and the other ventral to the chaetae. Notopodial chaetae are acicular. 
Neuropodial chaetae and chaetae in the anterior uniramous part are long hetero- 
gomph spinigers (Text-fig. 17, a). 

The large macrognaths each carry 12 teeth, six large and six small. There are 30 
dorsal and 12 ventral micrognaths. Papillae from the proboscis are illustrated in 
Text-fig. 17, b. About 150 pairs of chevrons have been counted, extending in two 
rows along almost the full length of the proboscis. Arwidsson reported 90 pairs in 
his original description, but it is not considered that the extra number present in 
the Accra specimen is sufficient to warrant specific distinction. In all other characters 
the specimens appear to be identical, and since the Accra specimen is almost twice 
as long as Arwidsson's the variation may be taken to indicate differences in growth 

Distribution. This species is known only from the tropical coast of West Africa. 



Fig. 17. Goniada multidentata. (a) Chaeta from the anterior uniramous region ; (b) 
group of papillae from the proboscis. 

Genus GONIADOPSIS Fauvel, 1928 

Body divided into three distinct regions ; (1) anterior, with uniramous parapodia 
and stout falcigerous chaetae ; (2) median, with uniramous parapodia and spinigerous 
chaetae ; (3) posterior, with biramous parapodia, notopodial acicular chaetae and 
long spinigerous neuropodial chaeta. Papillae on the proboscis are numerous, of one 
or few kinds. 

Goniadopsis incerta Fauvel, 1932 

Goniada (Goniadopsis) incerta Fauvel, 1932, pp. 122-123, pi- 4» n g s - 1-10, from Akyab, Burma. 
Goniadopsis incerta, Day, 1953, p. 430, from the estuary of the Zwartkops River, Cape Province, 
South Africa. 

Locality, (i) Apam shore, 4.1V.50 (1), 35 mm. long. 

The anterior region consists of 33 segments. Parapodia have three ligules, two 
anterior, finger shaped, of equal length, one posterior, broad and triangular. One 
of the stout falcigerous chaetae is shown in Text-fig. 18, a. Ventral cirri are short 
and blunt anteriorly, but elongate in the posterior segments of this region. 



The median region consists of 37 segments. Parapodial ligules are similar to those 
in the anterior region. One of the compound spinigers is shown in Text-fig. 18, b. 
Ventral cirri are very long, dorsal cirri are short. 

The posterior biramous portion makes up the remainder of the body of in seg- 
ments. There are two notopodial ligules, one posterior ; neuropodial ligules are 

Fig. 18. 

Goniadopsis incerta. (a) Falciger from the anterior region ; 
from the median region. 

(b) spiniger 

similar to those in the anterior region except that the posterior one is drawn out to 
a blunt point. Compound spinigers are as in the median region. 

There is one pair of small eye-spots at the base of the annulated prostomium. 

Distribution. This is the first record of this genus from the Atlantic Ocean. 

Genus OPHIOGLYCERA Verrill, 1885 

Body divided into three regions ; (1) anterior with uniramous parapodia, with 
spinigerous chaetae ; (2) median, in which notopodia with simple chaetae are 
gradually developed ; (3) posterior, with biramous parapodia. Proboscis long, 
terminated with a circlet of soft fleshy papillae, within which is a circlet of macrog- 
naths. Papillae on the proboscis are numerous of one or few kinds. 

Ophioglycera archeri sp. n. 

Holotype, B.M. (N.H.), Reg. No. 1953.3. 1. 648- 

Locality. Off Accra, Buchanan Survey, in 7-3 metres off Castle (1), complete, 
134 mm. long. The proboscis, which is not protruded, measures about 25 mm. long. 

The prostomium is without visible annulations or eye-spots. Eighteen large 
papillae form a crown at the end of the proboscis. The proboscideal teeth form a 

ZOOL. 3, 2. 9 



circlet, adjacent to the papillae, of two large separate macrognaths having between 
them groups of 18 and 25 micrognaths. The macrognaths have four teeth each, 
two large and two small, the micrognaths are bidentate. Papillae cover the surface 
of the proboscis and appear to be all alike (Text-fig. 20, b). 

On the first foot there is a single cirrus with no chaetae. Thereafter there are 27 
uniramous parapodia with spinigerous chaetae (Text-fig. 19, a), 49 segments in 
which notopodia are developed, but in which the rami are close together (Text-fig. 
19, b) and 140 biramous parapodia in which the rami are far apart (Text-fig. 20, a). 
Neuropodial chaetae are compound spinigers throughout, notopodial chaetae are 

In this specimen biramous parapodia are not gradually developed. At the 28th 
chae tiger the notopodial ramis appear fully developed with lobes and simple chaetae. 
A third division of the body, however, is recognizable at the 77th chaetiger where 
notopodia and neuropodia are much wider apart than they are anteriorly, and the 
chaetigerous ligules are much more foliaceous (compare Text-fig. 19, a and 19, b 
with Text-fig. 20, a). 

No diagnostic coloration is apparent in the preserved condition of the worm. 

No other species of Ophioglycera is known in which the change in form of the 
parapodia begins as far forward as in 0. archeri. In 0. foliocea the change begins at 
about the 35th chaetiger, but in 0. longicirrata, 0. gigantea, 0. eximia and 0. distorta 
it does not appear until the 55th or 59th, and, exceptionally, may be delayed to 
the 90th. 

<9 (9 

Fig. 19. Ophioglycera archeri, sp. n. Parapodia, (a) 13th foot, (b) 28th foot. 




Fig. 20. Ophioglyczva avchevi, sp. n. (a) Parapodium from a posterior foot of a mature 
$ ; (b) papilla from the proboscis. 


Family Eunicidae 

Body elongated, with the prostomium normally having one pair of palps and one 
to seven antennae. First two segments normally achaetous and apodous. Some- 
times one pair of tentacular cirri on the 2nd segment. Parapodia uniramous or 
sesquiramous. Dorsal cirrus, with or without branchiae, sometimes missing or 
rudimentary ; ventral cirrus often absent. Chaetae simple and /or compound, 
showing great variety of shape. Proboscis armed with a ventral mandible and dorsal 
maxillae. A membranous tube is secreted by some species. 

Key to Sub-families 

i. Prostomium reduced, without appendages (except in Augenaria Monro, with three 

minute antennae). No ventral cirri ; dorsal cirri rudimentary . . Lumbrinerinae. 

Prostomium with appendages, palps and antennae . . . . . . 2. 

2. No ventral cirri ; dorsal cirri foliaceous. Three antennae . . . Lysaretinae. 
Ventral cirri present ............ 3. 

3. Two antennae. Maxillae of two or four longitudinal series of very small and numerous 

pieces ............ Dorvilleinae. 1 

One to seven antennae. Maxillae of four or five pairs of pieces . . . . 4. 

4. Seven antennae, five occipitals mounted on ringed ceratophores and two ovate 

frontals ........... Onuphidinae. 

One to five antennae, ovate frontals absent ...... Eunicinae. 

Sub-family Eunicinae 

Key to Genera 

1. Branchiae present. Five antennae . . . . . . . . . 2. 

Branchiae absent ............ J. 

2. Tentacular cirri present .......... Eunice. 

Tentacular cirri absent .......... Marphysa. 

3. Three tentacles, tentacular cirri absent ....... Lysidice. 

One tentacle, tentacular cirri present ....... Nematonereis. 1 

Genus EUNICE Cuvier, 1817 

Five antennae. One pair of tentacular cirri on the second apodous segment. 
Dorsal cirri elongated, ventral cirri short or knob-like. Branchiae simple or pinnate. 
Parapodia sesquiramous, with acicular chaetae. Simple pectinate (comb-like) chaetae, 
and compound chaetae. Mandible of two pieces. Maxillae with a pair of forceps 
and two or three pairs of toothed plates, an unpaired left plate and sometimes 

There are manifold difficulties in identifying incomplete specimens of Eunice. The 
most important of these concerns the arrangement of those characters which appear 
on every parapodium, thus necessitating complete specimens for accurate diagnosis. 
A high degree of variability in the arrangement of branchiae, jaw plate dentition 
and extent of antennal annulation within any one species introduces further un- 
certainty. For these reasons all specific identifications of incomplete specimens are 

1 Not recorded from the Gold Coast. 


Key to Species 

i. Branchiae normally absent ; when present, simple, posterior, with one filament, 

sometimes 2-3 . . . . . . . . . . . E. gracilis. 

Branchiae, always present, pinnate . . . . . . . . .2. 

2. Subacicular hooks bidentate .......... 3. 

Subacicular hooks tridentate .......... 4. 

3. Branchiae occur only on the anterior part of the body beginning at about the 6th 

chae tiger ........... E. coccinea. 

Branchiae occur only on the middle and posterior part of the body, beginning at 

about the 26th chaetiger ........ E. filamentosa. 

4. Branchiae first present from about the 3rd chaetiger but absent over the greater part of 

the body from the middle backwards ...... E. vittata 

Branchiae first present from the 3rd to 7th chaetigers, and also present over the 

greater part of the posterior region . . . . . . . . . 5. 

5. Branchiae present on the last few chaetigers . . ■ . . E. antennata. 
Branchiae absent from the last few chaetigers . . . . . E. rubra. 

Eunice antennata Savigny, 1818 

Eunice antennata, Crossland, 1904, pp. 312-318, figs. 56-60. 
Eunice antennata, Hartman, 1944a, pp. 115-117, pi. 7, figs. 154-156. 

Localities, (i) Tenpobo shore, (1), anterior fragment, 28 mm. long. (2) Off 
Accra, Dredge in 14-6 m., shingle, 2 miles south out from the R. Densu (1), 70 mm. 
long, complete, and several fragments. 

The antennae are clearly annulated. Pectinate branchiae, in the dredged specimen, 
begin on the 7th chaetiger with 9 filaments ; on the 24th, 32nd and 52nd feet there 
are 12, 4 and 1 filament to each gill respectively. An increase to 3 filaments then 
takes place, which is maintained up to the last three segments on all of which there 
is a further reduction to one. Subacicular hooks are tridentate, compound chaetae, 
bidentate falcigers ; simple chaetae are capillary and pectinate. There are two 
pairs of anal cirri, one dorsal and long, the other ventral and short. 

Distribution. Cosmopolitan in tropical and subtropical areas. 

Eunice vittata (Delle Chiaje), 1829 

Eunice vittata, Augener, 1918, pp. 321-323. 

Eunice vittata, Fauvel, 1923, pp. 404-405, fig. 158, h-n. 

Eunice vittata, Hartman, 1944a, p. 118. 

Localities, (i) Tenpobo shore, (1), ant. piece. (2) Off Accra : (a) Stn. 35 (1), 
complete, 14 mm. long ; Stn. 47 (2), anterior pieces ; Stn. 69 (1) anterior piece ; 
Stn. 88 (1), complete, 8 mm. long; (b) Dredge in 14-6 m., shingle, 2 miles south out 
from the R. Densu (2), complete, 10 and 19 mm., long, and nine anterior pieces. 

In the specimen from Stn. 88 branchiae appear on the 3rd chaetiger with three 
filaments. A maximum development of 7-8 filaments occurs on the 38th foot, after 
which there is a reduction to one and finally branchiae are missing entirely from the 
posterior segments. Subacicular hooks, and chaetae, are similar to those in E. 

Distribution. Mediterranean, Atlantic, and Pacific. 


Eunice ftlamentosa Grube, 1857 

Eunice filamentosa, Augener, 1918, pp. 324-327, from Pram Pram. 
Eunice filamentosa, Hartman, 1944a, p. 107, pi. 6, figs. 123-126. 

Localities, (a) Axim shore, 13/14.^.49 (1), 40 mm. long, complete ; (b) Axim, 
Hospital Reef, 7.L51 (1), 200 mm. long, complete. 

In the specimen from Axim shore branchiae appear at the 29th chaetiger with one 
filament, gradually increase to a maximum of five and continue thus over the greater 
part of the body. Towards the pygidium there is a reduction to 3/4 filaments. The 
subacicular hooks which appear at the 23rd foot are strongly bidentate ; superior 
acicular have blunt, hammer-shaped ends. 

The specimen from the Hospital Reef has branchiae appearing at the 26th chaetiger, 
the maximum number of filaments is five and this condition prevails over much of 
the body. There is again a gradual reduction in filament number towards the tail, 
but in this specimen it is carried much further, the last three chaetigers having no 
branchiae. Fragments of a papyraceous tube are present. 

Distribution. Tropical West Africa, Eastern and Western Tropical America. 

Eunice coccinea Grube, 1878 

Eunice coccinea, Crossland, 1904, pp. 297-303, pi. 20, figs. 6, 7, text-figs. 46-51. 
Eunice coccinea, Fauvel, 1932, p. 136. 

Locality. Axim, Hospital Reef, 7.L51 (2), one complete, 42 mm., one incomplete, 
100 mm. long. 

Branchiae begin on the 6th and 7th parapodia. They attain a maximum of 10-12 
filaments in the anterior third and are missing throughout the rest of the body. 
Subacicular hooks appearing at the 25th and 28th feet, are bidentate. 

Distribution. Frequently recorded in the tropical regions of the Atlantic and 
Indo-Pacific, from the Gulf of Guinea to the Philippines. 

Eunice gracilis (Crossland), 1904 

Nicidion gracilis, Crossland, 1904, pp. 327-329, pi. 22, figs. 10-11, text-figs. 65-66. 
Eunice gracilis, Fauvel, 1932, pp. 140-141, text-fig. 20. 
Eunice gracilis, Day, 1949, p. 447. 

Localities, (i) Accra, (a) Christiansborg shore, 17.iii.49 (2), complete, 7 and 
n mm. long ; (b) Dredge 8 fathoms, shingle, 2 miles south of R. Densu (1), complete, 
20 mm. long. (2) Dixcove, shore (10), five complete, 15-42 mm. long. (3) Axim, (a) 
Shore, 13/14.^.49 (1), complete, 24 mm. long; (b) Lighthouse Reef, 8.L51 (1), 
complete, 20 mm. long; (c) Hospital Reef, 13.iv.49 (1), complete, 22 mm. long; 
7.L51 (10), seven complete, 10-18 mm. long. 

Simple chaetae are capillaries and comb-like ; compound chaetae are bidendate. 
Subacicular chaetae appear at about the 25th foot and are also bidendate ; there is 
one to each parapodium compared with two in E. cincta Kinberg, (see Hartman, 


1948, p. 80, on a re-examination of Kinberg's types). For this reason the synonymy 
proposed by Fauvel (1950) including E. gracilis as a synonym of E. cincta has not 
been accepted here. Antennae barely reach back to the 1st chaetiger ; tentacular 
cirri have frequently been lost. No branchiae are present ; anteriorly dorsal cirri 
are well-developed. 

There has been much discussion about the generic status of this species. Originally 
designated a species of Nicidion because of the lack of branchiae, Fauvel (1932) 
and Day (1949) have reported gills sometimes present on extreme posterior segments, 
and for this reason the species is regarded as a Eunice. 

Distribution. Indian Ocean — Zanzibar, Gulf of Manaar, Mergui peninsular — 
and Atlantic Ocean. St. Helena and the Gold Coast. 

Eunice rubra Grube, 1856 

Eunice rubra, Augener, 1918, pp. 319-321, from Pram Pram, and the Island of Rolas, in the 
Gulf of Guinea. 

This species has not been found in the present collections. The antennae are 
clearly annulated, branchiae begin at the 5th chaetiger and subacicular hooks are 
trident ate. 

Distribution. This species is common on the south-eastern coast of the U.S.A. 

Genus MARPHYSA Quatrefages, 1865 

Five antennae, two eyes. Tentacular cirri absent, dorsal cirri elongated ; 
ventral cirri short. Branchiae simple or pectinate. Dorsal chaetae simple capillaries. 
Ventral chaetae simple or compound, comb chaetae, and acicular chaetae also 
present. Mandible of two pieces. Maxillae with a pair of forceps, two pairs of 
toothed plates, an unpaired plate and sometimes paragnaths. 

Marphysa dartevellei Monro, 1936 

Marphysa dartevellei Monro, 1936&, pp. 246-248, 6 figs, from the Congo coast, Cape Malemba, 
85 km. north of Banana, from holes of Lithodomus. 

Localities, (i) Tenpobo shore, 5.U.50 (1), 325 mm. long, complete. (2) Accra, 
Christiansborg shore, 19.xi.49 (2), anterior pieces, 85 and 35 mm. long. (3) Axim, 
(a) Shore, 13/14.^.49 (2), anterior pieces, 26 and 35 mm. long ; (b) Hospital Reef, 
7.1.51 (1), 58 mm. long, complete. 

The antennae are unarticulated, equal in length, measuring only a little longer 
than the head ; between the laterals there are two black eyes. Dorsal cirri are smooth 
and slender decreasing in length posteriorly. Ventral glandular pads are present 
after the first few chaetigers. Ventral cirri are well developed stumps in the anterior 
feet, but posteriorly become little more than tubercles at the top of the glandular 
pads. Pectinate branchiae begin between the 32nd to 40th chaetigers. They have 
a maximum of four to five filaments and are present over the greater part of the 
middle body region. In the complete specimen from Tenpobo there are 86 


abranchiate segments posteriorly preceded by a large number of segments in which 
reduction to one filament has taken place. 

In the anterior parapodia compound chaetae are spinigers and posteriorly are 
falcigers. Monro's original specimens were not complete and his suggestion that 
falcigerous compound chaetae replace spinigers posteriorly is here substantiated. 
Simple capillary chaetae occur throughout the body, simple comb-chaetae are present 
in the middle body region but are often difficult to find posteriorly. 

There are two pairs of anal cirri, one pair elongated and dorsal, the other ventral 
and short. 

Distribution. This species is known only from the above records. 

Genus LYSIDICE Lamarck, 1818 

Three antennae. Tentacular cirri absent. Branchiae absent. Chaetae, simple 
capillaries, comb-like, compound falcigers and acicular. Mandible of two pieces. 
Maxillae with a pair of forceps, two toothed plates, an unpaired plate and paragnaths. 

Key to Species 

1 . Eyes round ; antennae small, never reaching as far as the anterior border of the 

prostomium ............ L. ninetta. 

2. Eyes reniform ; antennae reaching to the anterior border of the prostomium, some- 

times further . . . . . . . . . . . L. collaris. 

Lysidice ninetta Audouin & Milne-Edwards, 1834 

Lysidice ninetta, Augener, 191 8, pp. 362-364. 

Lysidice ninetta, Fauvel, 1923, pp. 411-412, fig. 162, a-g. 

Localities. Off Accra : Stn. 48 (1), 13 mm. long, complete ; Stn. 62 (1), 
6 mm. long, anterior piece ; from wood netted in 14 metres off Chorkor (1), 
21 mm. long, complete. 

For further details see below under L. collaris. 

Lysidice collaris, Grube, 1869 

Lysidice collaris, Crossland, 1903, pp. 143-144. 
Lysidice collaris, Fauvel, 1932, pp. 143-144. 
Lysidice collaris, Day, 195 1, pp. 39-40. 

Locality. Off Accra, Stn. 62 (1), 22 mm. long, complete. 

Differences, other than those noted in the above key, which serve to separate 
these two species are (a) a marked indentation on the anterior border of the prosto- 
mium compared with the almost entire border in L. ninetta, and (b) the greater 
development of the dorsal cirri in L. ninetta. 

The form of the jaw articles do not differ sufficiently to warrant emphasis. Day 
(195 1) has shown that this character is of value in differentiating between L. collaris 
and L. natalensis, particularly with respect to maxillae II. In L. collaris and L. 


ninetta, however, the former has four teeth on each article of maxillae II, the latter 
varying between 4-4 and 4-5. 

Chaetae are similar in both species. The simple types, capillaries and comb-like, 
are dorsal, the compound, with bidentate terminal articles, are ventral. In both 
species there are two pairs of anal cirri, a long dorsal pair and a short ventral pair. 

Distribution. L. ninetta is known extensively from, the European area in which 
L. collar is is unknown. There appears, however, to be a general overlap of the two 
species in the Pacific Panama, West African and Indian Ocean regions. L. collaris 
has been frequently recorded from the Indo-Pacific region, from Australia, China, 
Philippines, Japan, etc., in which areas no valid record of L. ninetta appears to have 
been made. The record of this latter species from New Zealand (Fyfe, 1952, p. 19) 
doubtfully accepted by Fauvel (1923, p. 411) and detailed by Augener (1924) as 
L. brevicornis Kinberg, is almost certainly L. collaris. 

Subfamily Onuphidinae 
Key to Genera 

1. Branchial filaments inserted spirally ........ Diopatra. 

2. Branchial filaments simple or pectinate ....... Onuphis. 

Genus DIOPATRA Audouin & Milne-Edwards, 1833 

Seven antennae, of which two are small front als and five occipitals borne on ringed 
ceratophores. One pair of clear, translucent, circular areas posterior to the inner 
laterals, which are probably sensory in function. One pair of tentacular cirri on 
the apodous peristomium. Dorsal cirri subulate ; ventral cirri subulate in a few 
anterior segments thence becoming pad-like. Pseudo-compound chaetae in anterior 
segments, otherwise with simple capillary, comb-like and acicular chaetae. Branchiae 
large with filaments inserted spirally. Mandible of two pieces. Maxillae with a pair 
of forceps, three pairs of toothed plates and one unpaired plate. Tube parchment- 
like with agglutinated material of varying types. 

Key to Species 

1. Tip of the shaft of the comb-chaetae continued as a central spike (Text-fig. 21, a). 

Tube as in Text-fig. 22, a . . . . . . . . D. musseraensis. 

2. Tip of the shaft of the comb-chaetae not so continued. Tube as in Text-fig. 22, b. 

D. neapolitana. 

Diopatra musseraensis Augener, 1918 

Diopatra musseraensis Augener, 1918, pp. 347-349, pi. 5, fig. 134 ; pi. 6, fig. 195 ; text-fig. 38, 
from Angola. 

Localities, (i) Off Accra, dredge, 14-6 metres, in shingle 2 miles south from the 
mouth of the R. Densu (1), incomplete, 40 mm. long. (2) Winneba shore, 22.xi.49 
(5), incomplete, 25-120 mm. long, with tubes measuring 160-240 mm. in length. 

Strongly spiralled branchiae are present on about 50 chae tigers. The anterior 
dorsal surface is uniformly coloured mauve. 



Chaetae are in well-defined groups. Prominent comb-chaetae (Text-fig. 21, a) are 
dorsal and anterior, they project almost as far as the stout winged capillaries which 
form a large group ventral to them. A bulbous parapodial cirrus projects beneath 
this latter group and divides the anterior set of pseudo-compound bristles from a 

Fig. 21. Comb-chaetae of (a) Diopatra musseraensis , x 500 ; (b) and (c) 
Diopatra neapolitana, x 750. 

small posterior group of capillaries. The pseudo-compound chaetae number four 
to a parapodium and have a darker brown terminal piece apparently grafted on to 
the light brown basal portion. This gives the effect of a possible articulating surface. 
Ventro-anteriorly to the ramal cirrus are two bidentate subacicular chaetae, which 
first appear at about the 17th parapodium. 
Distribution. Known only from the above records. 

Diopatra neapolitana Delia Chiaje, 1841 

Diopatra cuprea, Augener, 1918, pp. 350-354, text-fig. 39, from Elmina and Accra, Gold Coast, 

Diopatra neapolitana, Fauvel, 1923, pp. 419-420, fig. 166, a-h. 
Diopatra neapolitana, Fauvel, 1933, pp. 28-37, n g s - 3 _ 4- 

Localities. Off Accra : Stn. 6 (1), + tube ; Stn. 16 (1) ; Stn. 27 (1), + 
tubes ; Stn. 53 (1) ; Stn. 57 (34), -f tubes ; Stn. 58 (6), + tubes ; Stn. 59 (1) ; 
Stn. 61 (1), -f tubes ; Stn. 70 (6) ; Stn. 73 (6), + tubes, one specimen complete, 
30 mm. long ; Stn. no (1), with tube ; Stn. in (1), with tube ; Stn. 126 (1), com- 
plete, 27 m. long. Apart from the specimens from Stns. 73 and 126, all are less 



than 15 mm. in length and are anterior fragments. Buchanan Survey, 3-6 to iro 
metres off Accra (7), incomplete, up to 30 mm. in length. 

In the complete specimen from Stn. 126 branchiae begin on the 5th parapodium 
and are spiralled up to the 30th foot, after which they become branched with one 
or two filaments for 10 segments and then disappear. The anterior dorsal surface 
is skin-coloured, crossed, in some specimens, by lateral stripes of pale mauve. 



Fig. 22. Tubes of (a) Diopatra musseraensis, (b) Diopatra neapolitana. 

The form of the comb-chaetae in these specimens is illustrated in Text-figs. 21, b- 
c. These chaetae were exceedingly difficult to find and in no specimens were they 
as liberally distributed among the parapodia as in D. musseraensis. 

Fauvel (1932 & 1933) has drawn attention to the variation in form of the comb- 
chaetae in this species, and examination of the Gold Coast specimens supports his 
conclusions. In one specimen, for example, comb-chaetae with 6 and 14 teeth are 
present. For this reason Augener's record of 1918 of D. cuprea has been included as 
a synonym of D. neapolitana. With the exception of the scarcity of comb-chaetae 


in D. neapolitana the distribution of chaetae in the parapodia is very similar to that 
in D. musseraensis. 

Distribution. Atlantic Ocean, Mediterranean Sea, Red Sea, Indian and Pacific 

Genus ONUPHIS Audouin & Milne-Edwards, 1833 

Similar to Diopatra except that branchiae are not spirally coiled, but either simply 
branched or pectinate. 

Onuphis eremita Audouin & Milne-Edwards, 1833 

Onuphis landanaensis, Augener, 1918, pp. 339-343, pi. 5, figs. 135-138, pi. 7, fig. 197, text- 
fig- 36. 
Onuphis eremita, Fauvel, 1923, pp. 414-415, fig. 163, a-l. 

Localities, (i) Tenpobo shore, about 30 specimens with tubes, some complete, 
measuring 30 mm. in length ; associated with Audouinia punctata. (2) Off Accra : 
Stn. 57 (3), fragments, up to 18 mm. long ; Stn. 12 (1), anterior piece. 

Eyes are absent in this species. Branchiae begin on the first chaetiger with one 
filament ; after 10 to 20 segments there is a gradual increase to a maximum develop- 
ment of five filaments. There are tri- or bidentate hooded pseudo-compound chaetae 
in the first three to five feet. Other chaetae are simple limbate capillaries and 
comb-like ; acicular chaetae are bidentate. The specimens from Tenpobo are 
much smaller than those described by Fauvel. Tubes are of agglutinated sand 
grains with a few shell fragments. 

Distribution. Atlantic Ocean, Mediterranean Sea, Suez Canal, Madagascar, 
Ceylon, Madras, Mergui, Akyab, Caribbean Sea, and off the coast of South California 
down to Guatemala. 

Subfamily Lumbrinerinae 

Genus LUMBRINERIS Blainville, 1828 

Body long and cylindrical. Prostomium without palps, antennae or eyes. First 
two segments apodous and achaetous. Branchiae absent. Parapodia with two 
unequal ligules. Simple winged capillary chaetae and simple or compound hooks. 
Mandible bodice-like. Maxillae with four pairs of plates and two supports. 

Key to Species 

1 . Prostomium conical, pointed anteriorly ; no compound hooded hooks, simple hooks 

as Text-fig. 23, c . . . . . . . . . . L. impatiens. 

2. Prostomium rounded, blunt anteriorly with some compound hooded hooks (Text- 

fig. 23, a), simple hooks as in Text-fig. 23, b . . . . L. albifrons. 

Lumbrineris impatiens (Claparede), 1868 
Lumbriconereis impatiens, Fauvel, 1923, pp. 429-430, fig. 171, a-i. 

Localities, (i) Tenpobo shore, 4.ii-5o (1), anterior piece 13 mm. long. (2) Accra, 
Christiansborg shore (2), one complete, 78 mm. long ; one fragment, 20 mm. long. 



Parapodia with a short, anterior, rounded lobe, and a long, posterior, cirriform 
ligule. Anterior parapodia have simple winged chaetae and simple hooks with 
denticles and guard. The capillaries are gradually replaced in the middle and posterior 
region, leaving only two or three simple hooks (Text-fig. 23, c) to each chae tiger. 
The large maxillary plates II have four teeth each, III 2 +2 and IV 1 + 1. The 
maxillary supports are much longer than those described by Fauvel. 

Distribution. Atlantic Ocean, Mediterranean, Red Sea, Persian Gulf, Indian 

Fig. 23. 

Chaetae of Lumbrineris. (a) Compound hooded hook of L. albifvons, (b) simple 
hooded hook of L. albifvons, (c) simple hooded hook of L. impatiens. 

Lumbrineris alhifrons (Crossland), 1924 

Lumbriconereis albifvons Crossland, 1924, pp. 50-55, text-figs. 65-72, from the Cape Verde 

Locality. Off Accra, from wood netted off Chorkor 24. v. 51 (2), complete, 75 
and 22 mm. long. 

This species is characterized by the form of the compound chaetae (Text-fig. 23, a). 
These are, however, exceedingly difficult to find. The dental apparatus is illustrated 
in Text-fig. 24 and also serves to distinguish the species from closely related forms. 
Simple winged capillary chaetae are present with hooded hooks anteriorly, but are 
replaced by the latter posteriorly. As in L. impatiens, anterior parapodial lobes are 
short, posterior lobes are long. 

Distribution. The only authentic records of this species are those noted above ; 
the records by Monro (1933) from the Galapagos Islands and Pruvot (1930) from, 
New Caledonia are of doubtful significance. 

Subfamily Lysaretinae 

Apart from the characters noted above in the key, the following are of importance 
in this subfamily. Mandible of two pieces, maxillae with five pairs of symmetrical 
toothed plates and two long supports (Text-fig. 25). 


Fig. 24. Lumbrineris albifrons. Maxillae I to IV from the ventral surface. 

Genus HALLA Costa, 1844 

Prostomium oval with four eyes and three antennae on its posterior border. 
There are two anterior apodous and achaetous segments and no tentacular cirri. 
The dorsal cirri are large and foliaceous ; all chaetae are simple and acicular chaetae 
are present. Maxillae asymmetrical with two very long maxillary supports, (Text- 
fig. 250 

Halla parthenopeia (Delle Chiaje), 1828 

Holla parthenopeia, Fauvel, 1923, pp. 426-427, fig. 169, a-h, from Naples, Genes and Cadiz. 

Locality. Off Accra, Buchanan Survey, in 3-6 to ii'o metres depth (3), two 
anterior and one posterior pieces, between 85 and no mm. long. 

Collector's note : " This worm is brick-red in life, blue in preservative. Grows 
up to 25 in. (625 mm.) long." 

The specimens agree very well with Fauvel's description. The superior dorsal 
chaetae, however, are not as hispid as presented in his figure. Maxillary plates are 
illustrated in Text-fig. 25 and show the plates to have the following teeth : I 11-7, 
II 12-15, III 10-9, IV ii-ii, V 1-1. There is, however, variation in this character 
and in the other specimens here examined the formulae is : I n-8, II 12-13, III 
16-12 +, IV 17-12, V 1-1. Many of the teeth are frequently broken, making accurate 
counting difficult. A dorsal incision was made in the pharynx in order to prepare 
Text-fig. 25, and the sets of teeth turned outwards. It appears, therefore, as if 
maxillae I and II of the right hand side worked against each other. In effect this 
maxillae I, with the basal portion of II, bites against I of the left hand side during 

The large, foliaceous, dorsal cirri each bear an external notch basally, and the 
chaetae project laterally between the parapodial lobe and a secondary, anterior 
pedal flap. 


Distribution. This appears to be the first record of this species in the Atlantic 
Ocean, south of Cadiz. 

Family Orbiniidae 

Body divided into two regions : (a) thorax, short and flat with few segments, 
neuropodia flattened pads with or without stout bristles ; (b) abdomen, long and 
cylindrical, with numerous segments, neuropodia erect. Prostomium without ap- 

^=^j c§7^ 

Fig. 25. H alia par thenopeia. Maxillae from the dorsal surface. (See text, 
p. 120, for explanation.) 


pendages. Proboscis unarmed. Parapodia biramous with simple chaetae. Branchiae 
dorsal. Transverse rows of papillae often present on the ventral side of a number of 
thoracic segments. 

Key to Genera 

i. Thoracic neuropodia with vertical rows of pedal papillae .... Orbinia. 

Thoracic neuropodia without vertical rows of papillae or with only 2-3 . . . 2. 

2. Thoracic neuropodial chaetae entirely slender, pointed .... Haploscoloplos. 1 

Thoracic neuropodial chaetae include pointed and hook-like ones . . . Scoloplos. 

1 Not recorded from the Gold Coast. 

Genus ORBINIA Quatrefages, 1865 

Prostomium conical. One pair of lanceolate gills on each segment except a few 
anterior ones. Thoracic notopodia with serrated capillary chaetae; neuropodia pad- 
like with stout chaetae and papillae. Abdominal notopodia with capillary and forked 
chaetae ; neuropodia bilobed with capillary chaetae. 

Orbinia foetida var. linguistica (Orlandi), 1896 
A ricia foetida var. linguistica, Fauvel, 1927a, pp. 14-16, fig. 4, a-l. 

Locality. Off Accra, Buchanan Survey, in 3-6 to n-o metres (1), incomplete, 
26 mm. long. 

There are 22 segments in the thorax, branchiae begin on the 7th and ventral 
papillae appear on the 16th segment, with one pair and gradually increase to five 
pairs on segment 21 ; they are absent from segment 22. There are eight papillae 
on the posterior neuropodial lobe. Large lanceolate chaetae appear on the nth 
to the last thoracic chaetiger. Between rami on abdominal segments there are pro- 
nounced cirri. 

Distribution. This variety appears to have been recorded only from the Mediter- 
ranean Sea, but the stem species is widely known from the English Channel and 
Atlantic Ocean and has been collected at Madagascar. 

Genus SCOLOPLOS Blainville, 1820 

Similar to Orbinia, but without pedal papillae or with only very few, 1-3, and 
never with inter-ramal cirri. 

Key to Species 

1. With capillary chaetae in anterior thoracic neuropodia . . . . . . 2. 

Without capillary chaetae in anterior thoracic neuropodia ; with 23 thoracic seg- 
ments, with branchiae beginning at the 6th ; with 3 rows of hooks in thoracic 
neuropodia ........... 5. johnstonei. 1 

2. With pedal papillae on anterior thoracic neuropodia ; with 12 to 20 thoracic seg- 

ments, with branchiae beginning at the 9th to 17th. ... 5. armiger. 1 

Without pedal papillae on anterior thoracic neuropodia ..... J. 

3. Branchiae begin on anterior thoracic segments ..... 5. dubia sp. n. 
Branchiae begin on posterior thoracic or anterior abdominal segments 

5. madagascarensis. 
1 Not recorded from the Gold Coast. 


Scoloplos madagascarensis Fauvel, 1919 
Scoloplos madagascarensis Fauvel, 1919, pp. 433-434, pi. 17, figs. 81-86. 

Locality. Off Accra, Buchanan Survey, in 3-6 to n-o metres (2), anterior pieces, 
each 33 mm. long ; there are also two median pieces which may be detached portions 
of the anterior parts. 

In one of the anterior pieces the thorax is made up of 21 segments, of which the 
first nine have two rows of neuropodial hooks and the remainder one. Branchiae 
appear on the last thoracic segment, on which there is also one pedal papilla. In the 
other anterior piece there are 26 thoracic segments, all of which have one row of 
neuropodial hooks. Branchiae appear on the first abdominal segment. This piece 
is without pedal papillae. 

Both specimens have a few short forked chaetae in abdominal notopodia, one arm 
of the fork being about twice as long as the other. These chaetae are very easily 
broken and difficult to find. 

Fauvel (1919) originally described 5. madagascarensis with a thorax of 26-27 
segments and with branchiae appearing at the 21st or 22nd segments becoming 
fully developed at the 25th or 26th. It will be seen that the specimens from Accra 
agree in one or other of these characters, and it is suggested that the species may be 
more variable than has hitherto been indicated. 

Distribution. Previously recorded from Madagascar and Inhaca Island, Delgoa 
Bay, Transvaal (Day, 1951). 

Scoloplos dubia sp. n. 
Holotype ; B.M. (N.H.) Reg. No. 1953.3. 1 .1230. 

Locality. Off Accra, Buchanan Survey, in 3-6 to ii-o metres depth, complete, 
about 90 mm. long. 

The sharply pointed prostomium is without visible eyes. There are 22 thoracic 
segments. Thoracic neuropodial capillaries (Text-fig. 26, a) appear on every foot ; 
notopodial chaetae throughout the body are similar to these. Forked notopodial 
chaetae have not been seen. Thoracic neuropodial hooks are shown in Text-fig. 
26, b, they occur in three to four rows on each segment. In the posterior region of 
the abdomen the neuropodial aciculae become external and hooked (Text-fig. 26, c). 

Paratypes ; B.M. (N.H.) Reg. No. 1953.3. 1 . 1231-4, from the same locality as the holotype. 

These specimens are all incomplete, the longest measuring 65 mm. In all of them 
the branchiae appear at the 7th foot ; in two the thorax has 22 segments, in one 
21 and in the other 23. In other characters they are identical with the holotype. 

As suggested by its name the status of this new species is doubtful. It is un- 
doubtedly closely related to S. johnstonei, from which it is distinguished principally 
in having thoracic neuropodial capillaries and posterior hooked aciculae. Examina- 
tion of material from areas between the Gold Coast and the Cape of Good Hope (the 
distributional zone of 5. johnstonei) may clarify the relationship. 

ZOOL. 3, 2. IO 



Fig. 26. 


Scoloplos dubia sp. n. (a) Thoracic neuropodial chaeta, (b) thoracic neuropodial 
hook, (c) posterior abdominal parapodium. 

Family Spionidae 

Body not divided into distinct regions. Prostomium without antennae ; (some- 
times with frontal peaks) ; with palps and eyes. Biramous parapodia with foliaceous 
ventral and dorsal cirri ; dorsal branchiae on a number of segments. Chaetae simple 
capillaries and hooded hooks. 

Genus PRIONOSPIO Malmgren, 1867 

Prostomium without frontal peaks. Branchiae 3-1 1 pairs, often pinnate, confined 
to anterior segments. Simple capillaries and pluridentate hooded hooks are present. 

Prionospio pinnata Ehlers, 1901 

Prionospio africana, Augener, 1918, pp. 402-5, pi. 6, figs. 162-3, text-fig. 51. 
Prionospio pinnata, Fauvel, 1932, p. 173. 

Localities. Off Accra, Stn. 69 (2), incomplete, both 7 mm. in length ; Buchanan 
Survey, off Accra, 3-6 to n-o metres, depth (7), one complete, 55 mm. long, others 
incomplete 10-41 mm. long. 

Pinnate branchiae, if present, occur on the first three chaetigers only, frequently 
they have been accidentally lost. Several of the specimens from the Buchanan 


Survey still retain one of the long pair of palps, which extends from the posterior 
region of the prostomium to the 16th chaetiger. The palps are somewhat curved 
and grooved dorsally (Text-fig. 27). Their external margin is marked with mauve 

Fig. 27. Prionospio pinnata. Grooved palp from the left hand side of the prostomium. 
(Specimen from the Buchanan Survey.) 

vertical stripes. Four eyes are set in a trapezium on the prostomium, which is 
enclosed between two membranous wings. Within these wings the prostomium 
consists of a prominent longitudinal ridge. 
Distribution. Atlantic Ocean, Indian Ocean and Pacific Ocean. 

Family Cirratulidae 

Body short, cylindrical, with numerous segments ; head without appendages, with 
or without eyes. Proboscis smooth, unarmed. Dorsal palps or tentacular cirri on 
an anterior segment ; long slender branchiae inserted above the notopodia ; para- 
podia biramous ; chaetae simple capillaries and acicular hooks. 

Key to Genera 

1. Branchiae appearing on the same segment as the tentacular cirri . . . Cirratnlus. 

2. Branchiae appearing on a few segments in front of the one bearing tentacular cirri 


Genus AUDOUINIA Quatrefages, 1865 

First three segments achaetous. Branchiae occur on the first chaetigers to nearly 
the last ones. Of the three species of this genus recorded here, each is confined to a 
particular area of the Gold Coast. Thus A. punctata has been found only in the 
Tenpobo area ; A. tentaculata only on the Winneba shore, and A. filigera only in 
the Sekondi-Axim region. No overlap has been noted. 

Key to Species 

1 . Branchiae in the middle body region situated half the distance above the notopodium 

that the notopodium is above the neuropodium . . . A. tentaculata. 

Branchiae in the middle body region situated much more than half the distance 

above the notopodium than the notopodium is above the neuropodium . . 2. 


2. One hook in the neuropodium in the posterior part of the body. Branchiae in the 
middle body region situated, normally, much further than twice as far above the 
notopodium as the notopodium is above the neuropodium . . . . A . filigeva. 

Three or more hooks in the neuropodium in the posterior part of the body. 
Branchiae in the middle body region situated at least as far above the 
notopodium as the notopodium is above the neuropodium . . . A . punctata. 

Audouinia tentaculata (Montagu), 1808 

Cirratulus tentaculatus var. meridionalis, Augener, 1918, pp. 461-463, pi. 6, fig. 175 — 176. 
Audouinia tentaculata, Fauvel, 1927a, pp. 91-92, fig. 32, a-g. 

Locality. Winneba shore, 22.xi.49 (20), between 18-35 mm. long, and another 
ten specimens, badly mutilated, in association with tubes of Sabellaria eupomatoides 
from this area collected on the same day. 

The tentacular cirri appear between the 6th and 7th chaetigers. Typically there 
are two hooks in the neuropodia over the greater part of the body, though for some 
segments after the 12th foot there may be three to four. 

Distribution. Common in the European zone from the North Sea, Irish Sea, 
English Channel and the Atlantic Ocean. Known also from German South- West 
Africa (Augener, 1918) and the Cape (Day, 1951). 

Audouinia filiger a (Delle Chiaje), 1841 

Audouinia filigeva, Fauvel, 1927a, pp. 92-93, fig. 32, h-m. 
Audouinia filigeva, Fauvel, 1932, p. 178. 

Localities, (i) Sekondi, high level plateau, 21.ii.49 (4), complete, up to 20 mm. 
long. (2) Dixcove, shore (12), mutilated, 20-40 mm. long. (3) Princes Town, shore, 
15.iv.49 (4), complete, 20-25 mm. long. (4) Axim, Hospital Reef (3), 10-12 mm. 

The tentacular cirri occur between the 4th and 5th chaetigers. After the appearance 
of two neuropodial hooks at the 12th foot, which persist for a small number of seg- 
ments, there is a reduction to one, which condition is maintained up to the end of 
the body. 

Distribution. Well known from the Atlantic Ocean, Persian Gulf, and the Indian 
and Pacific Oceans. 

Audouinia punctata (Grube), 1858 

Civvatulus punctatus, Augener, 1918, pp. 465-467. 
Audouinia punctata, Day, 1951, p. 47. 

Locality. Tenpobo shore, approximately 370 specimens, measuring up to 20 mm. 
long were collected here during February, 1950. 

The tentacular cirri are situated between the 3rd and 4th chaetigers. There are 
normally three hooks in the neuropodium over the greater part of the body, but 
anteriorly (i.e. just after the 12th parapodia) there may be a reduction to two and 
posteriorly an increase to four. Colour in spirit grey-black, flecked with black, but 
considerable areas of white may be present ; white to grey tentacular cirri with 


black lateral stripes ; gills grey black with sometimes an orange tip ; young forms 

Distribution. This species is known from the Atlantic, south of Florida to the 
Cape of Good Hope, and from the Natal coast. 

Genus CIRRATULUS Lamarck, 1818 
Similar to Audouinia except as in above Key. 

Cirratulus filiformis Keferstein, 1862 
Cirratulus filiformis Fauvel 1927a, pp. 94-95, fig. 33, h. 

Locality. Off Accra, Buchanan Survey, in 3-6 to n-o metres (5), all incomplete, 
the largest piece measuring 38-0 mm. long. 

The prostomium is sharply pointed, and has no eyes. On the 1st chaetiger there 
is one pair of long and filiform branchiae and these are present up to the end of 
the body. All chaetae are simple capillaries, there are no hooks present. Anteriorly 
segments are narrow ; posteriorly they are long. 

Distribution. This species is well known from West European waters and has 
been reported from Senegal by Fauvel (1902). 

Family Flabelligeridae 

Prostomium and peristomium with eyes, palps and branchiae, completely re- 
tracted into the oral aperture, made visible only by dissection. Chaetae of anterior 
segments long, directed forwards to form a cephalic cage. Parapodia biramous and 
normally without distinct processes. Notopodial chaetae simple ; neuropodial 
chaetae sigmoid or hooked or compound with a sickle-shaped terminal piece. 

Genus STYLARIOIDES Delle Chiaje, 1828 

Body elongated, with numerous papillae on its surface. Prostomium with two 
stout palps. Branchiae filiform, on a peduncle, retractile. Notopodial chaetae 
simple capillaries, annulated ; neuropodia with simple capillaries and sometimes 
with strong hooks. 

Key to Species 

1. Neuropodial chaetae long and slender throughout . . . .5. scutigeroides. 
Some neuropodial chaetae hooks . . . . . . . . . 2. 

2. Neuropodial hooks with bidentate tips ...... 5. arenosus. 

Neuropodial hooks with entire tips ....... 5. tropicus. 

Stylarioides scutigeroides Augener, 1918 
Stylarioides scutigeroides Augener, 191 8, pp. 444-447, pi. 6, figs. 155 and 185, text-fig. 66. 

Localities. Off Accra : Stn. 55 (1) ; Stn. 69 (4) ; Buchanan Survey, in 3*6 to 
ii- o metres (5), only four of these specimens are complete, measuring between 
15 and 30 mm. in length. 


The complete body consists of 70 parapodia. The 1st chaetiger is telescoped into 
the 2nd, the 2nd into the 3rd and the 3rd into the 4th. Thereafter the segments are 
separated only by intersegmental grooves anteriorly and ridges posteriorly. 

Chaetae of the first parapodia vary between 2-5 to 4-5 mm. in length (Text-fig. 28, 
a and b) and are directed forward, forming the characteristic cephalic cage of the 
Flabelligeridae. Chaetae of the second parapodia are between 1 and 2-5 mm. long 
and are also directed anteriorly. Thereafter the chaetae decrease in size and project 
laterally. Notopodia and neuropodia in the first and second parapodia are close 
together, elsewhere they are far apart. The anterior region of the body is swollen, 
posteriorly it narrows, so that in one specimen the width across the 12th segment is 
2-5 mm. and across the 30th, 0-5 mm. Except for a few posterior segments the body 
is heavily encrusted with sand grains. 

All chaetae are long and slender ; notopodial bristles are illustrated in Text-fig. 
28, d, neuropodial bristles in Text-fig. 28, c. Papillae are present all over the body 
but are hidden to superficial view by the heavy encrustation of sand. On the first 
and second parapodia a single finger-shaped papilla, which may be bi- or trifid 
terminally, is associated with each group of chaetae, and there are sometimes one 
or two interramal in position. On the 3rd and 4th chaetigers simple finger-shaped 
papillae are present between the rami, encircling the segments like coronets. There 
are 6-7 dorsally between the notopodia, 2-3 laterally between the notopodia and 
neuropodia and 6-7 ventrally between the neuropodia. After the 5th segment the 
papillae become smaller and more scattered in arrangement. Posteriorly they are 
less numerous. 

Distribution. 5. scutigeroides is known only from the Gold Coast, the French 
Congo and Cabinda. 

Stylarioides arenosus (Kinberg), 1867 

Pycnoderma fernandense, Augener, 1918, pp. 448-452, pi. 6, figs. 148, 182 ; pi. 7, figs. 237, 
238 ; text-fig. 67, from Saltpond, Gold Coast. 

This species has not been collected in the present survey. Stout neuropodial 
chaetae, with bidentate tips appear at the 5th chaetiger. The secondary tooth at 
the tip is easily broken and the chaetae have frequently been described as unidentate. 
Papillae run along the body in longitudinal rows between the rami. Normally there 
are between 10 to 12 to each segment, four ventral and four dorsal and one or two 
laterally. One of Augener's largest specimens measured 39 mm. long for 75 segments, 
but Hartman (1948) found one of Kinberg's types measured 60 mm. for 70 segments. 

Distribution. Around the coast of Africa from the Gold Coast to Natal. 

Stylarioides tropicus Augener, 191 8 

Stylarioides tropicus Augener, 1918, pp. 437-440, pi. 7, figs. 220, 221, text-fig. 63. 

This species has not been collected in the present survey. It appears to be charac- 
terized by strong hooks with entire tips which appear at the 4th chaetiger, otherwise 
Augener's*" description is difficult to follow. One of Augener's specimens measured 
24 mm. long fori 32 segments. 




Fig. 28. Stylariodes scutigeroides. Chaetae ; (a) and (b) of the cephalic cage 
(c) neuropodial ; (d) notopodial. 


Distribution. The species was collected from Liberia, the Gold Coast, Accra, 
French Congo, Cabinda and Angola. 

Family Opheliidae 

Body short, convex dorsally, with or without a gutter ventrally. Prostomium 
without appendages, frequently with cephalic eyes. Sometimes with lateral eye 
spots. Biramous parapodia with simple chaetae. Pygidium with papillae or some- 
times prolonged into an anal tube. 

Key to Genera 

i. With lateral branchiae .......... Armandia. 

2. Without lateral branchiae ......... Polyophthalmus . 

Genus ARMANDIA Filippi, 1861 

Body not divided into distinct regions ; a deep median and two lateral ventral 
grooves. Branchiae from the 2nd chaetiger to the end of the body. Pygidium with 
anal funnel fringed with papillae and a median cirrus. Lateral eye-spots. 

Armandia intermedia Fauvel, 1902 

Armandia intermedia Fauvel, 1902, pp. 86-89, figs. 29, 30, from Senegal. 
Armandia intermedia, Augener, 1918, pp. 424-425, from Senegal and Angola. 
Armandia intermedia, Day, 1949, p. 449, from St. Helena. 

Locality. Off Accra, Stn. 69 (1), 12 mm. long. 

There are 29 chaetigers, the 1st abranchiate, the next 25 branchiate (though some 
of the posterior branchiae have fallen off, the point of attachment to the superior 
edge of the parapodia can be seen), and the last three abranchiate. Thirteen pairs 
of lateral eye-spots are situated on the 7th to the 19th chaetigers. There are three 
cephalic, subcutaneous, eye-spots disposed in a triangle. The anal funnel is fringed 
with numerous papillae, but many have fallen off and the exact number cannot be 

Distribution. This species is only known through the above records. 

Genus POLYOPHTHALMUS Quatrefages, 1850 
Similar to Armandia but with no branchiae. 

Polyophthalmus pictus (Dujardin) 1839 
Polyophthalmus pictus, Fauvel, 1927a, pp. 137-138, fig. 48, l-o. 

Localities, (i) Tenpobo shore, 6.ii.5o (2), 6-8 mm. long ; (2), Lighthouse Reef, 
Axim, 8.L51 (1), 9 mm. long. 

There are 28 chaetigers with about 11 pairs of eye spots on the 6th to 16th seg- 
ments, but these are very difficult to see and may vary considerably in position. 


Distribution. English Channel, Atlantic, Mediterranean, and the Red Sea ; 
Pacific and Indian Oceans. 

Family Maldanidae 

Body of few segments some of which are exceptionally long. Prostomium without 
appendages, often with a rimmed cephalic plate and a median keel, on each side 
of which is a nuchal groove. Buccal segment achaetous. Parapodia biramous, 
notopodial chaetae capillaries, neuropodial chaetae uncinate ; no parapodial cirri. 
Anal segment with funnel and cirri or a flat plate. Inhabitants of cylindrical tubes 
which may be thin and covered with sand or thick and coated with mud. 

Genus MALDANE Grube, i860 

Cephalic keel arched, rim divided into three by two lateral notches. Anus dorsal 
with a slanting anal plate. Ante-anal segments achaetous. Neuropodial chaetae 
missing from the first segment. Notopodial chaetae of three kinds. Tube coated 
with mud. 

Maldane sarsi Malmgren, 1865 
Maldane sarsi, Fauvel, 1927a, pp. 197-199, fig. 69. 

Locality. Off Accra, Buchanan Survey, in 36-5 metres off Castle (1), complete, 
about 62 mm. long. 

Prostomium with the cephalic keel strongly arched and the rim smooth, but 
notched on each side. 

The nuchal grooves are short, curved and deep. There are 19 chaetigers and two 
achaetous ante-anal segments. Notopodial chaetae are of three types : (a) long 
capillaries, barbed along their entire length ; (b) limbate capillaries, slightly barbed 
at their tips ; (c) very short geniculate capillaries with a limb on their convex borders. 

Uncini with one very large tooth surmounted by 5-6 rows of smaller teeth ; 
beneath the large tooth there is a cluster of fine hairs. 

Distribution. Cosmopolitan. 

Family Oweniidae 

Cylindrical body of few segments having the prostomium fused with the buccal 
segment and being either devoid of appendages or terminating in a membranous 
lobe. Notopodial chaetae simple capillaries ; neuropodial chaetae uncinate, very 
small, with a bent hooked tip. Tube coated with sand and shell fragments. 

Genus OWENIA Delle Chiaje, 1841 

Prostomium terminating in a membranous branchial lobe. Buccal segment 
achaetous ; the first three chaetigers without uncini. Notopodial chaetae slender, 
slightly spinous. Pygidium bilobed. 


Owenia fusiformis Delle Chiaje, 1841 

Owenia fusiformis, Fauvel, 1927a, pp. 203-204, fig. 71, a-f. 
Owenia fusiformis, Augener, 19x8, pp. 492-493. 

Localities. Off Accra : (a) Stn. 5 (25) ; Stn. 11 (4) ; Stn. 32 (numerous) ; Stn. 
53 (24) ; Stn. 54 (numerous) ; Stn. 55 (50 +) ; Stn. 61 (7) ; Stn. 65 (numerous) ; 
Stn. 66 (numerous) ; Stn. 67 (36) ; Stn. 121 (2) ; Buchanan Survey, 3-6 to n-o 
metres (9) ; (b) Dredge haul off Chorkor, 1 .xi.50 (20). 

The animals vary in length between 40-60 mm., the tubes, many of which are 
empty, particularly from Stns. 55, 65 and 66, measure up to 88 mm. The thorax 
consists of the achaetous buccal segment and the first three chaetigers which have 
only notopodial capillaries ; there is no neuropodial component in this region. The 
remaining segments, which are all biramous, make up the abdomen, and are long 
in the anterior region, but posteriorly become progressively smaller. The tubes 
consist of agglutinated small shells and sand, the former orientated at right angles 
to the long axis, and are grey in colour. 

Distribution. Cosmopolitan. 

Family Sabellariidae 

Body divided into four regions, (a) an anterior region with opercular disc and 
stalk, (b) a thorax of two anterior and three or four parathoracic segments, (c) an 
abdomen with uncinigerous dorsal rami and ventral rami with capillaries, (d) a 
caudal region, unsegmented, achaetous and apodous. The prostomium is hidden 
between the two large opercular stalks which carry paleae in concentric rows. Two 
palps. Simple branchiae are carried on the thorax and abdomen. Tubes are laid 
down and normally form sandy reefs of varying extent ; occasionally species may 
be solitary. 

Genus SA BELLA RIA Lamarck, 1818 

The operculum is short, with numerous filiform tentacles on the ventral side and 
provided with three visible rows of paleae. (The middle and inner rows probably 
represent a single row.) Thorax with two anterior segments, having only capillary 
chaetae, and three parathoracic segments with flattened paleae dorsally and similar, 
smaller chaetae, ventrally. 

Key to Species 

1. Animal small, less than 10 mm. in length when sexually mature ; tubes made up of 

small particles of sand . . . . . . . . .5. eupomatoides. 

Animals large, more than 15 mm. in length when sexually mature ; tubes made up 

of large and coarse sand grains . . . . . . . . . 2. 

2. Central tooth of outer paleae bluntly spatulate, entire or bifid 5. spinulosa var. intoshi. 
Central tooth of outer paleae with a barbed point , , 5. spinulosa var. alcocki. 



Sabellaria eupomatoides Augener, 1918 

Sabellaria eupomatoides Augener, 1918, pp. 503-504, pi. 7, figs. 187-190, text-fig. 83, from 

Localities. Numerous masses of the tubes of this species were collected from 
Winneba shore, Apam shore and Christiansborg shore at various times in 1949 ; 
specimens normally measure between 5-8 mm. in length. 

This species is characterized by its small size (a mature female from Winneba, 
collected 22.xi.49, measured 7 mm. long) and the form of the opercular paleae 
(Text-fig. 29). In some specimens several paleae of the median row project above 

Fig. 29. Sabellaria eupomatoides. 
(b) enlarged terminal piece of (a) 
row. (Specimen from Winneba.) 

Opercular paleae : (a) from the external row ; 
(c) from the outer row and (d) from the median 

the level of the others. There are nine pairs of simple gills from the 2nd thoracic 
segment backwards. 

S. eupomantoides was described by Augener in 1918 from specimens collected 
without their tubes, and has not been recorded since then except as above. 

Sabellaria spinulosa Leuckart, 1849 
Acicular chaetae are present on the dorsal face of the operculum, immediately 
posterior to the terminal crown of paleae. There is an inter-peduncular cirrus. 
Paleae in the external row have a barbed median tooth. 


Sabellaria spinulosa var intoshi Fauvel, 1914 
Sabellaria spinulosa var. intoshi Fauvel, 1927a, pp. 208-211, fig. 73^. 

Localities, (i) Tenpobo shore, about 20 specimens and tubes were collected in 
January and February, 1949, several were complete and measured up to 36 mm. 
long without the tail. (2) Apam (1), incomplete, 20 mm. long, and numerous tubes. 
(3) Dixcove shore (1), incomplete, 30 mm. long, with tube. (4) Axim Lighthouse 
Reef, 14.iv.49 (4), up to 38 mm. long, with tubes. 

This variety differs from the stem species in having the median tooth of paleae 
of the outer row with a spatulate tip. Occasionally this tip may be bifid. 

Distribution. English Channel and the Atlantic ocean (Gulf of Guinea). 

Sabellaria spinulosa var alcocki Gravier, 1906 
Sabellaria spinulosa var alcocki, Fauvel, 1927a, pp. 208-211, fig. 73, k-m. 

Locality : Axim, Hospital Reef, 7.L51 (1), 18 mm. long, with tube. 

Some of the paleae in the median row are always erect in this variety, otherwise 
it is similar to the stem species. 

Distribution. English Channel, North Sea, Atlantic Ocean, Indian Ocean and 
the Red Sea. 

Family Sternaspidae 

The form adopted in the family is aberrant. The body is swollen at both ends ; 
the prostomium small and devoid of appendages ; the first three segments have 
chaetae, the middle ones are achaetous, and posteriorly there is a ventral shield 
from the borders of which chaetae radiate. Branchiae are posterior to the shield. 

Genus STERN ASPIS Otto, 1820 
Branchiae filiform, set in two bundles. 

Sternaspis scutata (Ranzani) var. africana Augener, 1918 

Sternaspis fossor Stimpson var. africana Augener, 1918, pp. 608-613, text-fig. 109. 
Sternaspis scutata var. africana, Monro, 1930, pp. 179-180. 

Localities. Off Accra : Stn. 27 (42) ; Stn. 28 (2) ; Stn. 33 (6) ; Stn. 47 (2) ; 
Stn. 59 (1) ; Stn. 61 (8) ; Stn. 68 (14) ; Stn. 69 (19) ; Stn. 71 (3) ; Stn. 130 (1) ; 
Buchanan Survey, 3*6 to iro. metres (4). 

The specimens vary between 5 and 15 mm. in length, but these measurements must 
be treated with caution, for in many cases the anterior chaetigers have contracted 
into the succeeding segments. 

As Augener (1918) and Monro (1930) pointed out, the variety is distinguished, 
from the stem species by the greater number of chaetae (about 20 as opposed to 
10-12) which occur on the anterior segments, The chaetae are also more slender in 


the variety. These small differences seem to divide clearly the West African forms 
from the European and Antarctic examples, though further examination may 
indicate that the difference is merely ecological. Thus S. scutata has been reported 
from 68-1080 metres (Monro, 1930), from about 300 metres (Chamberlin, 1919), 
and S.fossor from 68-130 metres, (Fauvel, 1936) and from 120-130 metres, (Takahasi, 
1938), whereas S. scutata var. africana has not been reported beyond a depth of 
67 metres. Unfortunately many scattered records of the parent species do not give 
depth details or particulars of the morphological differences noted above, so that 
an accurate consideration of this issue is not possible. However, further collecting, 
at depth, off the Gold Coast may clarify the matter, and if it is confirmed that the 
morphological differences are non-genetic, but induced by the ecological conditions, 
the taxonomic treatment must be modified. If they prove to be genetic and asso- 
ciated with geography africana must rank as a subspecies. 

Distribution. 5. scutata var. africana is only known from off the W. African 
coast, from Senegal south to Angola. 

Family Ampharetidae 

Body divided into two regions, (a) thorax with notopodial capillaries and neuro- 
podial uncini, (b) abdomen with neuropodial uncini only. Tentacles retractile into 
the mouth. There are three or four pairs of branchiae on anterior segments and they 
may be subulate, pectinate or bi-pectinate. With or without groups of paleae. 

Key to Genera 

i. With paleae and without dorsal hooks behind the branchiae . . . . . 2. 

Without paleae and with dorsal hooks behind the branchiae .... Isolda. 

2. With one of the four pairs of branchiae pectinate ; 17 thoracic chaetigers Phyllamphicteis. 

With one of the four pairs of branchiae bi-pectinate ; 16 thoracic chaetigers Pterolysippe. 

Genus PHYLLAMPHICTEIS Augener, 1918 

With two fan-like groups of paleae ; thorax of 17 chaetigers with uncini appearing 
at the 4th. Abdomen of 15 chaetigers. Four pairs of branchiae, three subulate, one 

Phyllamphicteis collar ibranchis Augener, 1918 

Phyllamphicteis collaribranchis Augener, 1918, pp. 509-512, pi. 7, figs. 239-241, text-fig. 85, 
from Wappu on the Ivory Coast. 

Localities. Off Accra : Stn. 28 (1) ; Stn. 33 (1) ; Stn. 69 (1) ; Stn. 70 (1) ; 
Stn. 97 (4) ; Stn. in (1) ; several of these specimens are complete measuring up to 
20 mm. long. 

There are 8-10 paleae in each fan. Thoracic uncini have five teeth ; notopodial 
capillaries are slightly limbate. Both parapodial rami have small cirri on thorax 
and abdomen. The tubes are of fine dark grey mud with a few adhering shell particles. 

Four specimens collected in the Buchanan Survey, in 3-6 to iro metres off Accra 


and measuring between 20 and 30 mm. long, differ from the above in having two 
pairs of pectinate branchiae instead of one, though similar in other respects. It 
may be that they represent a different species, but it is possible that there is a 
considerable range of variation in this character within the species. Additional 
material and observations are needed to settle this point. 
Distribution. Only known from the above records. 

Genus PTEROLYSIPPE Augener, 1918 

With two groups of paleae, each like a fan. Thorax of 16 chaetigers with uncini 
appearing at the 4th. Four pairs of branchiae, 3 subulate and the other bi-pectinate. 

Pterolysippe bipennata Augener, 1918 

Pterolysippe bipennata Augener, 1918, pp. 512-514, plate 6, figs. 173 & 174, text fig. 86, from 
Whydah, Dahomey. 

Localities. Off Accra: Stn. 59 (1), with tubes; Stn. 133 (1) ; Buchanan Survey, 
in 3-6 to ii- metres (4), and in 41-86 metres off the Lagoon (1) ; several of these 
specimens are complete and measure up to 18 mm. long. 

The prominent bi-pectinate gill is coloured with numerous small purple marks. 
There are 15 abdominal segments. Thoracic uncini have 5-6 teeth, notopodial 
chaetae are limbate. Branchiae become easily detached in this species and it is 
rare to find a specimen with the complete set. No cirri are visible on the parapodia. 
Tubes are thick, of fine grey-black mud. 

Distribution. Only known through the above records. 

Genus ISOLD A Muller, 1858 

Without paleae. Number of thoracic and abdominal segments variable. Four 
pairs of branchiae, two subulate, and two feather-like. Two dorsal anterior hooks, 
immediately behind the branchiae. 

Isolda whydahensis Augener, 1918. 
Isolda whydahensis Augener, 1918, pp. 514-518, pi. 7, fig. 216, text-fig. 87. 

Locality. Off Accra, Buchanan Survey, in 5-5 metres off the breakwater (i), 
incomplete, length 15 mm. 

There are 16 thoracic chaetigers the first three of which are very difficult to see. 
They lie on the prostomial collar and in the specimen examined the chaetae had 
been broken off. Neuropodial uncini appear at the 5th chaetiger, are short and have 
five teeth. Thoracic capillaries are limbate. The transverse membrane which, with 
the prostomial collar, encloses the branchiae and dorsal hooks, is smooth. Both of 
the inner pairs of branchiae are feather-like, the outer pairs are subulate. 

Distribution. I. whydahensis is known also from Whydah, Dahomey, the French 
Congo and Angola. 


Family Terebellidae 

Body with thorax having notopodial capillary chaetae on all chaetigers and 
neuropodial uncini on some, and abdomen with neuropodial uncinionly. Prostomium 
with non-retractile tentacles. Branchiae, when present, normally on the anterior 
segments. In the thorax there are ventral glandular scutes. The tubes are coated 
with sand. 

Key to Genera 

1. Without branchiae ........... Amaea 

With branchiae ............ 2 

2. With a single quadripartite pectinate gill ...... Terebellides 

With more than one gill ........... 3 

3. Thorax with more than 17 chaetigers ....... Streblosoma 

Thorax with 17 chaetigers or less ......... 4 

4. Uncini pectinate ........... Loimia 

Uncini avicular ............ Pista 

Genus PISTA Malmgren 1865 

Thorax with 15-17 chaetigers ; eye-spots may be present. One, two or three pairs 
of branchiae. Notopodial chaetae smooth ; neuropodial uncini appear in the thorax 
on the 2nd chaetiger. 

Pista grubei Augener, 1918 
Pista grubei, Augener 1918, pp. 542-5, pi. 7, fig. 252-3, text-fig. 92. 

Localities. Off Accra: Stn. 28 (1) ; Stn. 61 (1) ; Stn. 69 (1) ; Stn. 131 (1). 

None of the specimens is complete, they vary between 25 and 47 mm. in length. 

There are 17 thoracic chaetigers, of which the first has only simple notopodial 
chaetae. In the next six segments there are also neuropodial groups of uncini in 
single rows, all the hooks lying in the same direction. The remaining 10 thoracic 
chaetigers, however, have the uncini lying alternately head to tail in a single row. 
The uncini are avicular in shape and all have a projecting basal process. 

The gills are in two pairs, consisting of a long basal trunk supporting a mass of 
whorled branching filaments. They are dorsal, anterior to the first chaetiger. 

Distribution. Known also from Senegal, Ivory Coast, and French Congo. 

Genus TEREBELLIDES Sars, 1835 

With a single gill divided into four pectinate arms. Notopodial chaetae long, 
tapering and winged. Uncini of two types, the thoracic elongated, acicular, the 
abdominal pectinate. 

Terebellides stroemi Sars, 1835 

Terebellides stroemi, Fauvel, 1927a, pp. 291-2, fig. 100, i-g. 
Terebellides stroemi, Fauvel, 1932, pp. 234-5. 

Localities. Off Accra : Stn. 28 (4) ; Stn. 33 (2) ; Stn. 59 (4) ; Stn. 73 (2) ; 
Stn. 133 (2). Most of these specimens are complete, measuring between 10 and 
30 mm. long. Buchanan Survey, 36-5 m. off Castle (1), 35 mm. in length. 


Body rarely with more than 60 segments, the first 18 being thoracic, with the gill 
as above. Segments 3 to 6 have their anterior border free. Notopodial chaetae are 
simple and begin on segment 3 ; neuropodial uncini appear at the 6th chaetiger and 
are long, geniculate, acidular hooks ; the remaining thoracic segments also have 
long uncini, but these end in blunt tips surmounted by small denticles. The 
abdominal uncini are avicular with a short base and transverse rows of teeth above 
the main fang. 

Distribution. Mediterranean Sea, Atlantic Ocean, Indian and Pacific Oceans. 

Genus STREBLOSOMA Sars, 1872 

Body with a large number of segments ; often with eyes ; with two or three pairs 
of branchiae (sometimes five?) on the 1st, 2nd, or 3rd chaetigers. Notopodial 
capillary bristles appear on the 2nd segment (1st branchiate). Neuropodial uncini 
appear on the 5th segment (4th chaetiger). 

Streblosoma persica (Fauvel), 1908 

Gvymaea persica, Fauvel, 1911, pp. 419-421, pi. 20, figs. 35-43. 

Pseudothelepus nyanganus, Augener, 1918, pp. 552-555, pi. 7, figs. 257, 258, text-fig. 96. 

Streblosoma persica, Fauvel, 1950, p. 384. 

Localities, (i) Tenpobo shore, about 100 specimens collected in January, 1949, 
and February, 195 1, measuring between 20-50 mm. in length. (2) Apam shore, 
16.ii.49 (12). (3) Dixcove shore, 7 . ii . 5 1 (12). (4) Axim, 13/14.^.49 (2). 

The species is characterized by (a) a semi-circle of eye-spots on the dorsal surface 
of the tentacular disc and (b) the shape of the uncini, which have a large basal tooth, 
paired smaller median teeth and above these a single smaller tooth with several tiny 
ones on either side of it. 

Notopodial chaetae occur on almost all segments. There are three pairs of bran- 
chiae, formed of numerous filaments, one pair on each side of the first three chaetigers. 
The first pair are based on transverse pads which form semi-circles around the anterior 
aspects of the notopodia of the first chaetiger. The remaining two are based on pads 
situated wholly on the dorsal side of the notopodia. 

Distribution. The species has been recorded from the Persian Gulf, the Gulf of 
Manaar, the Island of St. Thomas in the Gulf of Guinea, Angola and Senegal. 

Genus LOIMIA Malmgren, 1865 

Body with 17 chaetigers and three pairs of aborescent gills. Anterior segments 
have lateral lobes. Notopodial chaetae winged. Uncini pectinate, opposed back-to- 
back in double rows from the 7th to the 17th thoracic chaetiger. 

Loimia medusa (Savigny), 1822 
Loimia medusa Augener, 1918, pp. 539-541, text-fig. 91. 
Lattice fauvelii Day, 1934, PP- 7 I- 73> fig* J 4» a ~d- 

Locality. Off Accra, Buchanan Survey, in 36-5 metres off the Castle (1), an 
incomplete specimen of 34 chaetigers measuring 28 mm. in length. 


The three pairs of gills, of which the first is very much the largest, all lie anterior 
to the first chae tiger. The uncini have 6 teeth in both thorax and abdomen. 

Distribution. The species has been recorded from the English Channel, the 
Atlantic, Indian and Pacific Oceans, the Red Sea and the Persian Gulf. 

Genus AMAEA Malmgren, 1865 

Prostomium large, with many tentacles. Branchiae and eye-spots absent. Thorax 
of few segments, with notopodial chaetae. Abdomen consisting of an anterior 
achaetous portion and a posterior region with slender uncini. 

Amaea accraensis Augener, 1918 
Amaea accraensis Augener, 191 8, pp. 561-562, pi. 7, fig. 246, text-fig. 98, from Accra. 

This species has not been found in the present survey. The thorax has 11 chae tigers 
on the left hand side and 13 on the right. Notopodial chaetae are barbed capillaries. 
The achaetous abdominal region is about two-thirds as long as the thorax. There 
are 26-27 chaetigers posteriorly on the abdomen, bearing hooks with slightly bent 
tips. Augener's specimen measured 18 mm. 

Distribution. This species is only known from the type specimen. 

Family Sabellidae 

Body divided into two regions : (a) thorax, with few segments, having notopodial 
capillaries and neuropodial uncini, and (b) abdomen, normally with numerous 
segments having notopodial uncini and neuropodial capillaries. Ventral glandular 
shields divided by a longitudinal groove. First chaetiger with an entire, or notched 
collar. Branchiae encircling the mouth. Tubes of mucous, or membranaceous or 
horny, material. 

Key to Genera 

1. Thoracic neuropodia with a single row of avicular uncini, pickaxe-shaped chaetae 

absent .............. 2. 

Thoracic neuropodia with a row of avicular hooks and a row of pickaxe-shaped 

chaetae (fig. 30, a) . . . . . . . . . . Potamilla. 

2. Branchiae without dorsal sty lodes ........ Laonome. 

Branchiae with dorsal sty lodes ......... Dasychone. 

Genus LAONOME Malmgren, 1865 
Two symmetrical, semi-circular, branchial lobes not in the form of a spiral. 
Branchial filaments without stylodes. No pickaxe-shaped chaetae. 

Laonome punctur at a (Augener), 1918 

Demonax puncturatus Augener, 1918, pp. 576-580, pi. 6, fig. 170, 171, text-fig. 102. 
Euratella puncturata Monro, 1930, p. 203. 

Localities, (i) Accra : considerable quantities of tubes were collected from 
Christiansborg shore on 15.1.49, 14.ii.49, and 17. hi. 49. (2) Apam, 16.ii.49, 

ZOOL. 3, 2. II 


numerous specimens and tubes. (3) Axim shore, 13/14.^.49, numerous specimens 
but no tubes. 

The prominent prostomial collar opens dorsally and forms two triangular lappets 
ventrally. There are between 10 and 12 branchial filaments which, in most specimens, 
bear at least one distinct pigment band. Pigment spots are also present on the 
collar and between the rami of thoracic and abdominal feet. No clear division is 
discernible between pigmented and non-pigmented forms since specimens from all 
localities show the above pattern, but one separate and large collection from 
Christiansborg shore (15.L49) is completely without colour. The outside edge of 
the filaments is thickened in places suggesting rudimentary stylodes, but they are 
much less well developed than in the genus Dasychone. The branchial membrane 
reaches along one-sixth of the length of the branchial filaments ; the two palps are 
foliaceous and a little longer than the membrane. 

There are 4-5 thoracic chaetigers and about 100 abdominal segments. Neuropodial 
uncini appear at the 2nd foot. Thoracic capillaries are of two kinds, short with a 
prominent limbate termination (not spatulate) and long with a slightly limbate end ; 
thoracic uncini are short, avicular. Abdominal capillaries and uncini are identical 
with thoracic bristles. 

Augener (1918) placed this species in the genus Demonax Kinberg, which, however, 
has pickaxe-shaped chaetae in the thoracic neuropodia ; Monro, (1930) placed it 
in the genus Euratella Chamberlin, which has no collar and only one type of thoracic 
capillary. It is here considered that the species is referable to Laonome. 

Genus DASYCHONE Sars, G.O., 1861 

Both branchial lobes equal, not in the form of a spiral. Stylodes on the branchial 
filaments. Prominent collar. Pickaxe-shaped chaetae absent. 

Dasychone lucullana (Delle Chiaje), 1828 
Dasychone lucullana, Fauvel, 1927a, pp. 320-321, fig. no, m-s. 

Localities. Off Accra: Stn. 35 (1), 8 mm. long; Stn. 73 (1), 15 mm. long; 
Stn. 131 (1), 16 mm. long, all specimens complete. 

The two branchial lobes are each made up of 13-14 filaments which carry slender, 
colourless stylodes dorsally. Along the filaments there are numerous violet pigment 
spots, which, when the branchiae are closed, give the effect of successive girdles of 
colour along the lobes. A small branchial membrane unites the bases of the gills ; 
there are two prominent canaliculate palps. 

The body, other than the branchiate portion, is uniformly brown, with a small 
violet pigment spot between the parapodial rami. There are 8 thoracic and between 
30 and 40 abdominal segments. Thoracic uncini which appear at the 2nd chaetiger, 
are avicular with slight extension at the base ; thoracic capillaries are limbate, 
short and long. Chaetae in the abdominal region are similar to thoracic types except 
that long capillaries, only slightly limbate, appear in far posterior segments. 

Distribution. Mediterranean and Adriatic Seas and from Senegal. 


Genus POTAMILLA Malmgren, 1865 

Branchial lobes symmetrical, not in a spiral, without stylodes. With a prominent, 
lobed collar. Thoracic notopodial capillaries limbate and spatulate, neuropodial 
chaetae avicular uncini and pickaxe-shaped. Abdominal notopodial uncini avicular 
and neuropodial capillaries limbate. 

Potamilla casamancensis Fauvel, 1902 

Potamilla casamancensis Fauvel, 1902, pp. 101-105, fig. 46-55, from the estuary of the River 
Casamance, Senegal. 

Localities. Off Accra : Stn. 12 (1), complete, 16 mm. long ; Buchanan Survey 
in 11 metres off Lighthouse (2), complete, 14 and 24 mm. long ; Buchanan Survey 
in 5-5 metres off the Lagoon (1), incomplete, 8 mm. long. 

There is an indentation in the collar dorsally, forming a deep groove which con- 
tinues down the dorsal surface of the thorax ; ventrally there is only a very slight 
break in the anterior border of the collar. 

Ten to sixteen filaments make up each branchial lobe ; the filaments are without 
eye-spots. There are 5-7 thoracic chaetigers ; uncini and pickaxe-shaped chaetae 
appear at the second. The limbate capillaries on the first segment are in a longi- 
tudinal row, thereafter they are lateral. Spatulate capillaries appear at the second 
chaetiger and are normally few in number. Limbate capillaries are present in all 
segments, both thoracic and abdominal ; spatulate chaetae have not been seen in 
the abdomen. Uncini in the thorax are avicular with long shafts ; in the abdomen 
they are short. 

Distribution. P. casamancensis is only known through the above records. 

Family Serpulidae 

Body divided into two regions ; [a) thorax, with few segments, having notopodial 
capillaries and neuropodial uncini, and (b) abdomen, with numerous segments, 
having notopodial uncini and neuropodial capillaries. First segment with a collar. 
With a thoracic membrane. Branchiae form a funnel around the mouth. Usually 
with an operculum. Tube calcareous. 

Key to Genera 

1. Body symmetrical ............ 2. 

Body assymetrical. Tube spirally coiled ....... Spirorbis. 

2. Without an operculum .......... Salmacina. 

With an operculum ............. 3. 

3. Chaetae of the first chaetiger bayonet-shaped with two basal stumps . . . 4. 
Chaetae of the first chaetiger without basal stumps . . . . . . 5. 

4. Operculum simple, funnel-shaped ........ Serpula. 

Operculum compound, with a central crown of spikes ..... Hydroides. 

5. Operculum globular, transparent ........ Apomatus. 

Operculum horny, with a cylindrical or conical cap , Vermiliopsis. 

I 4 2 


Genus SALMACINA Claparede, 1868 

Without an operculum and with few branchiae which are frequently bent distally. 
Uncini pectinate with a large inferior tooth. 

Salmacina incrustans Claparede, 1868 
Salmacina incrustans, Fauvel, 1927a, pp. 378-380, fig. 129, /. 

Localities. Numerous specimens were collected from : (1) Tenpobo shore, on 
rocks, 4-ii.5o ; (2) Winneba rock face, sheltered side, 22.xi.49, forming compact 
masses about the Polyzoan, Waterispora cucullata ; (3) Apam shore, 16.ii.49 ; (4) 
Axim hospital reef, 49, L.W.M., some specimens were present on a tube of 
Sabellaria spinulosa var. alcocki. 

The majority of these specimens measure between 2 and 3 mm. Tubes are white, 
cylindrical in shape, and very fragile. 

Fig. 30. Chaetae of (a) Potamilla casamancensis , pickaxe-shaped bristle from a thoracic 
neuropodium, x 350 (stn. 49) ; (b) Salmacina incrustans, a notopodial chaeta from 
the first thoracic segment x 200, (Winneba) ; (c) and (d) Hydroides arnoldi, thoracic 
and abdominal uncini respectively, x 400 (e), (/) and {g) Apomatus similis : (e) 
A pornatus -type chaeta from a posterior thoracic segment, x 100, (/) limbate chaeta 
from a thoracic segment, (g) scythe-like chaeta from a median abdominal segment 
X 150; (h) Serpula vermicularis , trumpet-shaped abdominal chaeta, x 200 (Stn. 115) ; 
(;') Vermiliopsis multicristata, geniculate abdominal chaeta, x 200. 

There are four pairs of gill filaments and seven thoracic chaetigers. A notopodial 
chaeta from the first of these is shown in Text-fig. 30, b. This form is characteristic 
for the species. 

Distribution. Mediterranean Sea and Atlantic Ocean. 


Genus HYDROIDES Gunnerus, 1768 

Chaetae of the first chaetiger bayonet-shaped, with two humps at the base of the 
blade. Thoracic chaetae limbate ; abdominal chaetae trumpet shaped. Uncini in 
thorax and abdomen with coarse teeth. Operculum funnel-shaped, with radii 
ending bluntly or in fine points, and with a crown of horny spines rising from its 

Pixell (1913) suggested that Hydroides should include those species with opercular 
spines having lateral processes and that species devoid of these lateral processes 
should be included within the genus EupomcUus. This division is not adopted in this 

Key to Species 

1. Opercular spines all alike ........... 2. 

Opercular spines not all alike ........ H. spinosus. 

2. Opercular spines with lateral processes . . . . . . H. norvegica. 

Opercular spines without lateral processes . . . . . . . H. arnoldi. 

Hydroides norvegica Gunnerus, 1768 
Hydroides norvegica, Fauvel, 1927a, pp. 356-7, fig. 122, i-o. 

Locality. Scraped off copper sheathing of boat and body, Gold Coast, (50). 

The species is immediately recognizable by its opercular structures. The basal 
disc is soft and white, bearing marginally between 20 and 30 crenulations. Fre- 
quently the distal horny disc is missing but, when present, it carries 13 to 15 large 
spines with smaller lateral ones. A small vertical spine is situated in the centre of 
the opercular disc. The operculum itself is a modified branchial filament, the ventral- 
most of one side or the other ; whichever it may be, and there is no constancy, the 
other is always broken off. 

Chaetae of the first of the seven thoracic chaetigers have two humps below the 
terminal point and below the humps there are a few fine denticulations. Thoracic 
uncini have seven teeth and abdominal uncini five. 

Distribution. Cosmopolitan. 

Hydroides spinosus (Pixell), 1913 
Eupomatus spinosus Pixell, 1913, p. 78, pi. 8, fig. 5. 

Localities, (i) Tenpobo, No. 1 Reef, 21.xi.49 (1), complete, 15 mm. long. 
(2) Off Accra : (a) Trawled 2 miles west of Densu River, 2. hi. 49 (1), incomplete, 
10 mm. long ; (b) Stn 99 (3), one complete, 10 mm. long, on tubes of Turritella 
annulata Kiener ; Stn. 127 (1), complete, 11 mm. long, on a tube of T. annulata. 

These specimens differ only slightly from the original description (Pixell, 1913). 
Thus the basal disc has frequently more than 32 long, pointed teeth and the number 
of tall strong spines on the upper disc, is not constantly 11 but varies, some speci- 
mens having 8, 9 or 10. The shape of all the spines except one is identical, they each 
have a terminal hook pointing outwards, a near-terminal one pointing down wards 


and inwards and an inner median near the base. The exceptional one is longer than 
the others, curves over them, and has only a basal hook. 

Distribution. H. spinosus has previously been recorded from the Suez Canal 
area and the Cape of Good Hope. 

Hydroides arnoldi Augener, 191 8 

Hydroides arnoldi Augener, 1918, pp. 595-598, text-fig. 107, pi. 6, figs. 151, 152, from Togo, 
Dahomey, and the Island of Annobon. 

Locality. Axim shore, 49 (3), incomplete, 4-11 mm. long. 

There are seven opercular spines which have an inner basal hook but are devoid 
of other processes. The largest, and possibly oldest, specimen has almost straight 
spines with blunt ends (Augener, 1918, pi. 6, fig. 152), but the smallest, and possibly 
youngest, has spines ending in a curved tip. It is suggested that normal wear on 
the operculum has produced these different shapes and the species may be expected 
to vary in this character even more than is noted here. 

Thoracic and abdominal uncini are illustrated in Text-fig. 30, c and d. 

Distribution. H. arnoldi is only known from the above records. 

Genus APOMATUS Philippi, 1844 

With a globular transparent operculum. Branchial filaments with eye-spots and 
short connective membrane. No special notopodial chaetae on the first thoracic 
segment, which is without neuropodia, but with characteristic Apomatus-type 
bristles on the last segments (Text-fig. 30, e). 

Apomatus similis Marion and Bobretzky, 1875 
Apomatus similis, Fauvel, 1927a, pp. 385-387, fig. 131 k—p. 

Locality Off Accra, Stn. 115 (10), with tubes, on the Polyzoan Steganoporella 
buskii Harmer. The specimens measure up to 6 mm. in length. 

The branchial lobes are made up of 10 to 16 filaments each. The second dorsal 
filament carries a globular transparent operculum distally, but it is otherwise un- 
modified and combines a branchiate and operculate function. This represents a 
primitive stage in the modification of a branchial filament to an operculum. Com- 
plete modification is found in the majority of Serpulidae, but there is no trace of it 
in Filograna and Protula. 

There are seven thoracic and well over 60 abdominal segments. The majority of 
thoracic chaetae are limbate (Text-fig. 30, /), but in the last 3-4 segments of the 
thorax characteristic Apomatus-type bristles appear (Text-fig. 30, e). Anterior 
abdominal segments are achaetous. Neuropodia in the middle of the abdomen have 
very small scythe-like chaetae (Text-fig. 30, g), but in four posterior segments long 
capillaries are present. Abdominal and thoracic uncini are alike, with numerous 
small teeth and a long basal one. 

Tubes are white and semi-cylindrical, with slight lateral crenulations. 

Distribution. English Channel, Mediterranean Sea, Atlantic Ocean, 


Genus SERPULA Linnaeus, 1758 

Chaetae of the first thoracic segment bayonet-shaped, with two humps at the 
base of the blade. Operculum funnel-shaped with radii ending bluntly along the 
margin. Thoracic chaetae limbate, abdominal chaetae trumpet-shaped. Uncini 
with few, but stout teeth. 

Serpula vermicularis Linnaeus, 1767 
Sevpula vermicularis, Fauvel 1927a, pp. 351-352, fig. 120, a-q. 

Localities, (i) Tenpobo reef, 12.xi.49 (2), incomplete, 10 and 20 mm. in length, 
without tubes. (2) Off Accra, Stn. 115 (3), complete, up to 20 mm. in length. 

These specimens appear to be smaller than is normal for the species (Fauvel, 
1927a), but those from Stn. 115 were taken from tubes completely attached to the 
polyzoan, Steganoporella buskii, none of the tubes growing free, and they may be 
juveniles. The tubes are pink in colour, and triangular in shape with a prominent 
dorsal keel. 

There are 16-20 branchial filaments in each lobe united by a short basal membrane. 
The first dorsal filament of one side is completely adapted as an operculum, with a 
funnel-shaped termination having a crenulated surface. The first dorsal filament on 
the opposite side to that bearing the operculum is partially modified ; it is similar 
to the opercular filament, but is smaller and lacks the funnel-like termination. 

There are seven thoracic chaetigers the first without neuropodia. Apart from the 
bayonet-shaped chaetae with two basal humps on the first thoracic segment, 
notopodial-chaetae are unmodified capillaries. Abdominal neuropodial chaetae are 
trumpet -shaped (Text-fig. 30, h). 

Distribution. Cosmopolitan. 

Genus VERMILIOPSIS St. Joseph, 1906 

Operculum a horny cylindrical or conical cap. No special chaetae on the first 
chaetiger, which is without neuropodia. Apomatus-type chaetae in posterior thoracic 
segments. Abdominal chaetae geniculate (Text-fig. 30, j). 

Key to Species 

1. With seven thoracic chaetigers ........ V. multicristata. 

2. With nine thoracic chaetigers ....... V. prampramiana. 

Vermiliopsis multicristata (Philippi), 1844 
Vermiliopsis multicristata, -Fauvel, 1927a, pp. 365-366, fig. 125, k-s. 

Locality. Off Accra, Stn. 115 (6), some complete, up to 10 mm. long, taken from 
tubes attached to the polyzoan Steganoporella buskii. 

This species is characterized by its tube bearing 5-6 longitudinal ridges. There 
are about 10 branchial filaments to each lobe. The operculum is horny and cylindrical 



to conical in shape. In the thorax there are seven chaetigers and in the abdomen 
upwards of 50. The majority of thoracic chaetae are limbate, but in posterior 
segments a few Apomatus-type bristles appear. Abdominal chaetae are short and 
geniculate (Text-fig. 30, j) anteriorly, but posteriorly long capillaries appear. 

Distribution. Mediterranean Sea and Atlantic Ocean — Madeira and the Azores. 

Vermiliopsis prampramiana Augener, 1918 

Vevmiliopsis prampramiana Augener, 1918, pp. 603-604, pi. 7, fig. 256, from Pram Pram. 

This species is only known through this record, and is the only species of Vermiliop- 
sis known with nine thoracic chaetigers. Augener's description is otherwise in- 
complete, and it is difficult to assess the species relationship with other members 
of the genus or family. 

Genus SPIRORBIS Daudin, 1800 

Body assymetrical, with less than five thoracic segments. Opercular peduncle 
without pinnules. Tubes spirally coiled, dextral or sinistral. 


Key to Sub-genera 

Tube dextral ........ ... 2 

Tube sinistral ....... 


With 4 thoracic segments ..... 


With 3 thoracic segments ..... 


With 4 thoracic segments ..... 


With 3 thoracic segments ..... 


Chaetae of the first thoracic chaetiger limbate 


Chaetae of the first chaetiger with a crenulated wing 

. (Laespira) 

This genus was, surprisingly, absent from the collections, but it is almost certainly 
present in the Fauna, and, for this reason the above key has been included, (from 
Fauvel, 19270, pp. 389-391). Augener (1918, pp. 607-608), reported Spirorbis sp. 
from the " Cape Coast Castle", probably Christiansborg Castle, but his description 
does not permit further identification. When, and if, species are collected further 
reference to Fauvel should be made. 


Arwidsson, J. 1898. Studien iiber die Familien Glyceridae und Goniadidae. Bergens Mus. 
Aarborg, 1898, No. 11 : 1-70, 4 pis. 

191 1. Die Maldaniden. Wiss. Ergeb. der Schwed. Siidpolar-Exped, 1901-3, 6, No. 6: 

1-44, 2 pis. 

Augener, H. 1913. Polychaeta Errantia. Die Fauna Sudwest-Austt -aliens, 4 : 65-304, 
2 pis., 42 text-figs. 

1918. Polychaeta. Beitrage zur Kenntniss des Meeresfaunas West-Afrikas, 2:67-625, 

6 pis. 

1924. Polychaeta von Neuseeland. I, Errantia. Medd. naturh. Foren. Kobenhaven, 

75 : 241-441, 11 figs. 


Baird, W. 1864. Description of a new species of Annelide belonging to the family Amphino- 
midae. Trans. Linn. Soc. London, 24 : 449-450, pi. 45. 

Bergstrom, E. 1914. Zur Systematik der Polychaeten familie Phyllodociden. Zool. 
Bidrag. Uppsala, 3 : 37-224, 81 text-figs. 

Chamberlin, R. V. 1919. The Annelida Polychaeta. Mem. Mus. Comp. zool. Harvard, 
48 : 1-5 14, pis. 1-80. 

Claparede, E. 1868. Les Annelides Chetopodes du Golfe de Naples. Mem. Soc. Phys. 
Geneve, 19 : 313-584, 16 pis. 

Crossland, C. 1903. On the marine fauna of Zanzibar and British East Africa, from collec- 
tions made by Cyril Crossland in the years 1901 and 1902. Polychaeta, Pt. 2, Eunicidae. 
Proc. Zool. Soc. London, 2 : 129-144, pis. 14 and 15, text-figs. 12-15. 

1904. The marine fauna of Zanzibar and British East Africa from collections made by 

Cyril Crossland in the years 1901 and 1902. Polychaeta, Pt. 3. Ibid., 1 : 287-330, 
pis. 20 to 22, text-figs. 43-66. 

1924. Polychaeta of Tropical East Africa, the Red Sea and Cape Verde Islands collected 

by Cyril Crossland, and of the Maldive Archipelago collected by Professor Stanley Gardiner, 
M.A., F.R.S. Ibid. Pt. 1 : 1-106, text-figs. 1-126. 

Day, J. H. 1934. O n a collection of South African Polychaeta, with a Catalogue of the 
species recorded from South Africa, Angola, Mosambique and Madagascar. J. Linn. 
Soc. London, Zool. 39 : 15-82, 16 text-figs. 

1949. Polychaeta of St. Helena. Ibid. 41 : 434-451, 4 text-figs. 

1 95 1. The Polychaet Fauna of South Africa. Pt. 1 : The Intertidal and Estuarine 

Polychaeta of Natal and Mozambique. Ann. Natal Mus. 12 : 1-67, 8 text-figs. 

1953- The Polychaet Fauna of South Africa. Pt. 2 : Errant Species from Cape Shores 

and Estuaries. Ibid. 12 : 397-441, 1 map. 6 text-figs. 

Eiilers, E. 1887. Reports of the annelids of the dredging expeditions of the U.S. coast 
survey steamer "Blake." Mem. Mus. Comp. Zool. Harvard. 15 : 1-335, °° plates. 

Fauvel, P. 1902. Ann61ides Polyene tes de la Casamance (Senegal, Africa). Bull. Soc. 
Linn. Normandie, 5th series, 5 : 59-105, figs. 1-55. 

191 1. Annelides Polychetes du Golfe Persique. Arch. Zool. Exper. gen. Paris. (5), 

6 : 353-439, pis. XIX-XXI. 

1914- Annelides Polychetes de San Thome (Golfe de Guinee). Ibid. 54: 105-155, pis. 


1 91 9. Annelides Polychetes de Madagascar, de Djibouti, et du Golfe Persique. Ibid. 

58 : 315-473, Pis- 15-17- 

1923. Polychetes errantes. Faune de France, Paris 5 : 488, 1-181 text-figs. 

1927a. Polychetes sedentaires. Ibid. 16 : 1-494, 152 text-figs. 

19276. Contribution a l'etude de Faune du Cameroun. Polychaeta. Faun. Colon. 

franc, 1 : 5^3-533, 2 figs. 

1932. Annelida Polychaeta of the Indian Museum, Calcutta. Mem. Indian Museum 

Calcutta 12 : 1-262, 9 pis., 40 text-figs. 

1933- Annelides Polychetes du Golfe du Pei Tcheu Ly de la collection du Musee Hoang 

ho Pai ho. Publ. Mus. Hoang. ho Pai do de Tien Tsin, 15 : 1-67, 6 text-figs. 

1936. Remarques sur les Nereidiens, Nereis succinea Leuckart et Nereis lamellosa Ehlers. 

Bull. Soc. Zool. France, 61 : 307-314. 

1950. Contribution a la Faune des Annelides Polychetes du S6negal. Bull. Inst, francais 

d'Afrique noire, 12 (2) : pp. 335-394. 

Fyfe, M. L. 1952. List of New Zealand Polychaetes. Bull. New Zealand Dept. Sci. Industrial 

Research, 105 : 1—38. 
Gravier, C. 1909. Ann61ides polychetes recueillis a Payta (Perou). Arch. zool. exp. gen. 

Paris, 10 : 617-659. 
Hartman, O. 1939. Polychaetous Annelids. Pt. 1 : Aphroditidae to Pisionidae. Allan 

Hancock Pacific Exped. 7 : 1-156, 28 pis. 

1944a. Polychaetous Annelids. Pt. 5 : Eunicea. Ibid. 10 : 1-238, 18 plates. 

zool. 3, 2. 12 


Hartman, O. 19446. Polychaetous Annelids, Part 6, Ibid., 10 : 311-389, pis. 27-42. 

1948. The Marine Annelids erected by Kinberg with notes on some other Types in the 

Swedish State Museum. Ark. Zool. Stockholm, Band 42A, no. 1 : 1-137, pis. 1-18. 

1950. Polychaetous Annelids, Goniadidae, Glyceridae, Nephtyidae. Allan Hancock 

Pacific Expd. 15 : 1-181, 19 plates, 3 text-figs. 

1 95 1. The Littoral Marine Annelids of the Gulf of Mexico. Publ. Inst. Marine Sci. 

Univ. Texas 2 (1) : 7-124, 27 pis. 

Kinberg, J. 1856. Nya slagen och arter af Annelider. Fork. Oefv. Vet. Akad. Stockholm 
12 : 381-388. 

1857-1910. Kongliga Svenska Fregatten Eugenics Resa omkring Jovden under C. A. 

Virgin 1851-53. Zool. 3, Annulater, Uppsala-Stockholm, 78 pp., 29 plates. 

McIntosh, W. C. 1885. Report on the Annelida Polychaeta collected by H.M.S. " Chal- 
lenger " during the years 1873-6. " Challenger " Rep. Zoo. 12 : 1-554, pl s - I_ 55 an d 

1903. Marine Annelids (Polychaeta) of South Africa, I. Marine Investigations in South 

Africa, 3 : 17-56, 4 pis. 

1924. Preliminary note on a collection of South African polychaetes. Report Union 

South Africa Fish Mar. Biol. Survey, Cape Town, 3, No. 2, 1 p. 

Monro. C. C. A. 1930. Polychaeta Worms. " Discovery " Reports, 2 : 1-222, 91 text-figs. 

1933- The Polychaeta Errantia collected by Dr. C. Crossland at Colon, in the Panama 

Region, and the Galapagos Islands during the Expedition of the S.Y. " St. George ". 
Proc. Zool. Soc. London, 1933, (1) : 1-96, 36 text-figs. 

1936a. Polychaete Worms 2. " Discovery " Reports, 12 : 59-198, 34 figs. 

19366. Notes on some Polychaeta from the Congo Coast. Rev. Zool. Bot. Afr. 28 : 245- 

248, 1 fig. 

Pixell, H. L. M. 191 3. Polychaeta of the Indian Ocean, together with some species from 

the Cape Verde Islands. The Serpulidae. Trans. Linn. Soc. London, 2nd. Ser., Zoology, 

16 : 69-92, pis. 8 and 9. 
Pruvot, G. 1930. Annelides polyene tes de Nouvelle-Caledonie recueillies par M. Francois. 

Arch. Zool. exp. gen. Paris, 70 : 1-94, 8 figs., 3 pis. 
Quatrefages, M. A. de. 1 865. Histoire des Anneles, 1, Paris, 588 pp. 
Takahasi, K. 1938. Polychaeta, collected by the " Misago " during the Zoological Survey 

around the Izu Peninsula. Sci. Rep. Tokyo Bunrika Daigaku, Sec. B, 3 (57) : 192-220, 

1 pi. 
Treadwell, A. L. 1900. The polychaetous Annelids of Porto Rico. Bull. U.S. Fish Comm. 

Washington, 20 : 18 1-2 10, 81 figs. 
Willey, A. 1904. Littoral Polychaeta from the Cape of Good Hope. Trans. Linn. Soc. 

London. Zool. 9 (2) : 255-268, pis. 13-14. 
1905. In Herdman. Roy. Soc. Report on Pearl Oyster Fisheries, Ceylon. Pt. 4 : 243- 

324, pis. 1-8. 

2 SEP 1955 


2 JUL 1955. 







ZOOLOGY Vol. 3 No. 3 

LONDON : 1955 




(Bingham Oceanographic Laboratory, Yale University) 

Pp. I51-167. 



ZOOLOGY Vol. 3 No. 3 

LONDON: 1955 

(NATURAL HISTORY), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical Series. 

Parts will appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

This paper is Vol. 3, No. 3 of the Zoological series. 


Issued June, 1955 Price Five Shillings 




In spite of the efforts of Robson (1929), which form a landmark in the history of 
octopodan taxonomy, the Octopus rugosus problem remains unsolved, an outstanding 
source of confusion obscuring the correct identification of small, rough, vulgaris-like 
specimens from all parts of the world. Some years ago the writer reviewed the western 
Atlantic forms which had been, or could be, assigned to Octopus rugosus. It was 
possible to show that no such species can be found in American waters and that 
western Atlantic specimens previously referred to it were either 0. vulgaris or, 
occasionally, belonged to one of the other Caribbean species (Pickford, 1945, 1946). 

It is believed that the problem can best be solved by a zoogeographical approach, 
pari passu with increasing knowledge of each local fauna to the point that makes 
the recognition of its component species a matter of certainty. Through the work 
of Voss (1950, 1 95 1, etc.) and the author we have now arrived at this state of know- 
ledge of the American east coast fauna. However, the situation in other parts of 
the world is less satisfactory. Octopodan taxonomy has always been hampered by 
the lack of adequate series of specimens collected from any one locality and this, 
together with the confusing effects of different states of preservation, has made 
recognition of the limits of variation within a single species all but impossible. 

In the present contribution an attempt has been made to clarify the status of 
those specimens from the eastern Atlantic and South Africa which Robson identified 
as 0. rugosus and which are now in the collections of the British Museum. A survey 
of bodily characteristics, irrespective of species assignation, is given in Table I. It 
will be seen that only 7 of the 22 specimens are sexually mature, 4 are well grown 
but immature, and n are juvenile. In a difficult problem of this nature it is 
obvious that no definite conclusions can be based on juvenile, nor even on sexually 
immature specimens, except in so far as they can reasonably be assigned to adult 
forms with well established characteristics. 

Octopus rugosus, as it is generally understood, is a species closely resembling 0. 
vulgaris but smaller in size ; the head is broader, the arms shorter, and the rough, 
finely granulated skin is darkly reticulated with black. None of these characters, 
taken singly, can be used with any degree of certainty for the separation of the two 
species. Taken together the resulting picture is indistinguishable from that of 
juvenile and immature specimens of 0. vulgaris, so that it may be conjectured that 
0. rugosus is merely the young of 0. vulgaris (Pickford, 1945). This point of view 
is that taken by the writer who believes that it is necessary to prove that an Atlantic 
specimen assigned to 0. rugosus is not a young 0. vulgaris before it can be considered 
to belong to any other species ; a rugosus-like specimen should be assigned to 0. 
zool. 3, 3. 13 


vulgaris unless there is conclusive evidence that it does not belong to this species. 
Robson, on the other hand, used the name 0. rugosus as a hold-all to receive those 
vulgaris-like specimens about whose identification he was uncertain and, as a result 
of this policy, it is not surprising to find that the majority of his eastern Atlantic 
specimens, as indicated above, are juvenile or immature. 

We may approach the problem from another angle. Bosc's Sepia rugosa came 
from Senegal and the type is lost. Lamarck (1799) gives no locality for the two 
specimens to which he gave the name 0. granulatus. He points out that this species 
may be merely a variety of 0. vulgaris and concludes that it is probably the same 
as Bosc's rugosus. The types are lost and, in the absence of evidence to the contrary, 
it appears advisable to relegate the name 0. granulatus to the synonymy of 0. 
vulgaris. A similar conclusion is probably justified in the case of 0. tuber culatus 
Blainville, the type of which is also lost. However, since the species is from Sicily, 
its status should be re-examined in connexion with a more thorough taxonomic 
study of the Mediterranean population. 

It must be pointed out that the original specimens of 0. rugosus and 0. tuber cu- 
latus, not to mention numerous other specimens referred by various authors to these 
species or to 0. granulatus, were reported from geographical areas in which 0. 
vulgaris is known to occur. Therefore, 0. rugosus (to confine the discussion to one 
of the three problematical forms) is not a geographical race or subspecies of 0. 
vulgaris. Either it is a distinct species or it is inseparable morphologically from the 
extremes of the 0. vulgaris population. This is an important consideration. If two 
species occur together in the same geographical area they must form two separate 
breeding populations. 

The search for 0. rugosus is therefore the search for a West African vulgaris-like 
sibling species which is distinct as. a breeding population from 0. vulgaris with 
which it is sympatric. As the result of previous investigations (Pickford, 1945, 1946) 
it can definitely be stated that 0. rugosus does not occur in the western Atlantic 
but the problem of Indo-Pacific forms that have been assigned to this species awaits 
further clarification. However, if the existence of a true 0. rugosus cannot be 
established in the eastern Atlantic it is highly improbable that anything recorded 
from Indo-Pacific waters could legitimately be assigned to this species. 

The study of the British Museum specimens reviewed below leads to the following 
conclusion : 

(1) Nearly all specimens from the eastern Atlantic are probably or certainly 
referable to 0. vulgaris. It is probable that the correct assignation of the juvenile 
specimens will always remain in doubt but 6 of the 7 adults present no features 
which would permit their separation from 0. vulgaris. 

(2) Among the series there is one juvenile specimen from Madeira that has a 
distinctive pattern of zebra bands on the arms. Although the characteristics (Table I, 
B.M., 1898. 5. 10.2) are similar to those of 0. vulgaris the correct assignation of this 
specimen must remain in doubt. 

(3) One specimen from the Cape Verde Islands (1889. 4. 24. 17), a mature male, 
certainly represents a distinct species on account of the length of the ligula which 
is quite outside the range for 0. vulgaris. A similar male and also a female that 


apparently belong to the same species have been described by Adam (1952) from 
the Cape Verde Islands as Octopus sp. The question arises whether these individuals 
should be arbitrarily assigned to 0. rugosus, with which they show some general 
agreement, or whether it would be preferable to designate them by a new name. 
The original description of 0. rugosus does not permit recognition with any degree 
of certainty. Bosc's figures (1792, Figs. 1 and 2), if correctly drawn, show an animal 
of 42 mm. mantle-length with moderately long arms (MAI 32) and an unusually 
shallow web (WDI 13-5) ; the suckers are of moderate size (SnI 9 • 5) but none are 
specially enlarged. In favour of assigning the Cape Verde specimens to this species 
is the geographical area from which it came, as well as the general appearance and 
bodily proportions, except for the shallow web. The only contrary evidence is the 
shallow web and the uncertainty whether Bosc's 0. rugosus was not, after all, just 
a young vulgaris. For the sake of clarity and to avoid the confusion that might 
result from the use of a name which is, by now, so clouded in obscurity, it has 
seemed preferable to propose a new name for this Cape Verde species. 

(4) The problem of the specific affiliation of the South African rugosus-like popu- 
lation remains unsettled. The individuals that have been examined, from the Cape 
Province and Natal, appear to belong to the same species and to represent a moderately 
uniform group. Robson admits (1929, p. 59) that true 0. vulgaris occurs in South 
African waters. The two males, from Port Elizabeth and from Natal respectively, 
have the characteristic minute ligula of 0. vulgaris}- Massy (1925) found the same 
feature in males which she identified as 0. granulatus in the collections of the Natal 
Museum (LLI 1-7 and 1-65 respectively, calculated from her measurements). 
There is a small but possibly significant detail regarding the repeated reversal of 
the direction of coiling of the horn in the spermatophore of one of the males (B.M., 
1890 .9.4.4), but our information concerning the range of variation in spermatophores 
of 0. vulgaris is inadequate to assess this feature. Also the females, with ripe ovaries, 
are unusually small for sexually mature females of 0. vulgaris ; but this again 
amounts to no more than a suggestion. Possibly the South African forms represent 
a geographical race or subspecies of 0. vulgaris with these slightly different features. 
Such a conclusion could only be confirmed by the statistical analysis of a long series 
of specimens. 

A detailed account of the specimens follows : 


English Channel 

(1) B.M., 1927. 2. 10. 2. Octopus rugosus (Bosc), Robson (1929). Plymouth. C.92. 

It was pointed out in a previous paper (Pickford, 1946) that this specimen is 
clearly a female of 0. vulgaris. I take this opportunity to publish the characteristics 
listed in Table I and to call attention to the following features : the arms are un- 

1 Attention must again be called to Robson's mistake in tabulating the ligula-length indices for 
O. vulgaris. It appears that in this one case he referred the length of the ligula to the mantle length 
instead of to the length of the hectocotylised arm, thus giving misleadingly high values (see Pickford, 



usually short and the web rather deep but the suckers and gills are normal ; there 
is a very conspicuous lateral seam round the mantle, no doubt an accident of pre- 
servation ; the ovary is quite small and the ova are minute. 

Table I. — Characteristics of specimens from the eastern Atlantic, and South Africa 
which were referred by Robson (1929) to the species Octopus rugosus (Bosc), in the 
collections of the British Museum* 


English Channel 

1927.2. 10.2 

$ imm. 










6* juv. 









1898.5. 10.2 

$ imm. 







ca. 8 







Canary Islands 






















Cape Verde Islands 


$ imm. 









6* juv. 








1889.4.24. 17 

6* mat. 









1889.4.24. 14 

$ imm. 







ca. 10 








West Africa 

1928.3. 17. 1 

£ mat. 







ca. 6-' 

1928.3. 17.2 









1928.3. 17.3 









1889.4.24. 15 



South Africa 


c£ mat. 








1924.9.9. 11 

$ mat. 


1924.9.9. 12 

$ mat. 



$ mat. 



$ mat. 


(28) J 







1889.4.24. 18 



* Two females from the " South Coast " could not be found (B.M., 1872 .2.3. 12-13). Three specimens 
(1865.5.23. 1) from an unknown locality omitted. Data based on new measurements unless otherwise 
indicated. Symbols as in Pickford (1945). 

t Data from Hoyle (1886). 

I Data from Massy (1927). 



(2) B.M., 1867.3.27.9. Octopus rugosus (Bosc), Robson (1929). Mediterranean, 
purch. of Mr. Cutter. C.104. 

This is a very young male, not a female as listed by Robson. As far as can be 
determined it is probably referable to 0. vulgaris. 


(3) B.M., 1898.5. 10.2. Octopus rugosus (Bosc), Robson (1929). Porto Santo, 
Madeira, H. S. Wellcome Esq. C.86. 

This specimen, an immature female, is remarkable for its coloration. There are 
light and dark zebra bands on the arms and it is very dark and blotchy on the 
back and head. The ovary is undeveloped but has moved to a ventral position and 
is possibly rather advanced for a vulgaris of this size ; the ova are minute. The 
jaws have been dissected out and the radula is mounted on a slide. The bodily 
proportions are similar to those of a typical vulagris but on account of the peculiar 
coloration I hesitate to refer the animal to this species. It will be necessary to 
await further collections from this region to establish the existence of a form with 
banded arms and, until such time, it may be considered to be merely an unusual 
specimen of 0. vulgaris. 

(4) B.M., 1912.12.31.118 and 119. Octopus rugosus (Bosc), Robson (1929). Pedro 
de Malha, Porto Santo, Madeira, 60 fathoms, R. Kirkpatrick Esq. C. 152-153. 

Two very young specimens with the characteristic rhomb of 4 papillae on the 
back commonly seen in juvenile specimens of 0. vulgaris ; probably correctly 
referred to this species. 

Canary Islands 

(5) B.M., 1954. 2. 25. 1-3. Octopus rugosus (Bosc), Robson (1929). Canaries, R. 
McAndrew. C. 100-102. 

Three very young specimens, probably all referable to 0. vulgaris. The two 
larger specimens each have the characteristic rhomb on the back. 

Cape Verde Islands 

(6) B.M., 1851.1.24.1. Octopus rugosus (Bosc), Robson (1929). St. Jagos, C. de 
Verd, Ch. Darwin Esq., No. 122. C. 85. 

An immature female, very dark, mottled and rough. The ovary is small and 
apical ; the ova minute. The jaws have been dissected out. The suckers are large, 
as in vulgaris, the arms moderately long and the web moderately deep. There seems 
to be no reason why this specimen should not be assigned to 0. vulgaris. 

(7) B.M., 1855.9.29.2. Octopus rugosus (Bosc), Robson (1929). St. Vincent, Mr. 
MacGillivray coll. C. 119. 


This specimen is an immature male. The head is rather warty, the body smooth. 
The total absence of a hectocotylus at a mantle-length of 31 mm. indicates that it 
belongs to a large species and there seems to be no reason why it should not be 
assigned to 0. vulgaris. 


(8) B.M., 1889. .4.24.14 Octopus occidentalis Hoyle (Steenstrup MS), Hoyle (1886) ; 
Octopus rugosus (Bosc), Robson (1929). Challenger Coll., Ascension, April 1878. 
C. 69. 

This specimen, an immature female, is the type of 0. occidentalis and was described 
by Hoyle (1886) in the Challenger Reports. In its present state it is very dark and 
hard. A comparison of its present measurements with those given by Hoyle (Table 
I) indicate that a very considerable shrinkage of the arms has taken place during 
sixty years of preservation (the specimen was examined by me in 1946) but the 
length and proportions of the mantle and head are little changed. The longest arm, 
the second left, now measures about 180 mm. whereas Hoyle gives a length of 
230 mm. No doubt his measurements, which represent the condition of the specimen 
when it was in a better state of preservation, should be accepted. 

Robson could find no reason why this specimen should not be assigned to 0. 
rugosus and I can see no reason why it should not be assigned to 0. vulgaris. 

West Africa 

(9) B.M., 1928.3. 17. 1. Octopus rugosus (Bosc), Robson (1929). Old Collection 
(Rich), West Africa. Bears manuscript name " Octopus rugosus var. longi- 
manus. Type." C. 260. 

This is a mature male, very dark in color. The bodily proportions are those of 
vulgaris (Table I) the ligula (Table II) is minute and typical of vulgaris. The gills 
are very damaged and the low estimate of the number of gill lamellae may be due 
to this condition. Spermatophores were removed from Needham's organ ; the 
length (9 mm.) gives an index of 37-5, which is typical of vulgaris. The horn is 
very slender and measures about 4 mm. in length ; there are 40 to 50 spiral turns, 
partly uncoiled ; only the distal extremity of the horn, proximal to the middle 
piece, is not coiled. This last mentioned feature is not characteristic of a well pre- 
served spermatophore of vulgaris but the partial uncoiling and poor state of preserva- 
tion may well account for such a condition (compare Adam, 1952, Fig. 53B). There 
seems to be no reason why this specimen should not be assigned to 0. vulgaris. 

(10) B.M., 1928.3. 17.2. Octopus rugosus (Bosc), Robson (1929). Sierra Leone, 
W. P. Lowe. Bears manuscript name " Octopus rugosus ? var. longimanus." 
C. 262. 

A juvenile specimen, rather warty, with 4 dorsal papillae in a rhomb on the back. 
Probably referable to 0. vulgaris. 

(11) B.M., 1928. 3. 17. 3. Octopus rugosus (Bosc), Robson (1929). Murray Town, 
Sierra Leone, W. P. Lowe. C. 371. 


This juvenile specimen is very dark and extremely hard and shrunken. It is 
very probably a young vulgaris. 

(12) B.M., 1889.4.24. 15. Octopus tuberculatus Blainville, Hoyle (1886) ; Octopus 
rugosus (Bosc.), Robson (1929). Challenger Coll., No. 122B, off Barra Grande, 
Lat 9 9' S., Long. 34 53' W., depth 32 fm, red mud, 10.ix.1873. C. 115a. 

This specimen was identified as 0. tuberculatus by Hoyle, but is not described. 
There are indications of the dorsal rhomb of papillae and the specimen is very 
probably referable to 0. vulgaris. 

South Africa 

(13) B.M., 1890.9.4.4. Octopus rugosus (Bosc), Robson (1929). Port Elizabeth, 
South Africa, H. S. Spencer. Bears manuscript label " Octopus granulatus 
Lam." C. 108. 

A mature male with all the characteristics of 0. vulgaris. The specimen is very 
hard but one spermatophore was recovered from Needham's organ and was found 
to be in a moderately good state of preservation. Its length (10-5 mm) gives an 
index that is typical of vulgaris (Table II). There are about 37 coils to the spiral 

Table II. — Characteristics of adult male specimens referable to Octopus vulgaris Lam* 

B.M. No. 

West Agrica 

1928.3. 17. 1 
South Africa 

1890.9.4.4 . 

1924.9.9. 11 

* Symbols as in Pickford (1945). 

horn, interrupted by two irregular regions of partial uncoiling. The direction of the 
spiral is reversed several times, as follows : 8 coils anticlockwise, about 6 irregular 
clockwise coils, an irregular break, 3 anticlockwise coils followed immediately by 5 
clockwise turns, an irregular break, 8 anticlockwise coils followed by 6 clockwise. 
This repeated reversal of the spiral is not a characteristic feature of vulgaris spermato- 
phores but there has been no large scale survey of spermatophore variation in this 
species and, in the absence of such data and of an adequate series of spermatophores 
from South African males, it seems unwise to attach any great significance to the 
condition. Apart from this feature the specimen appears to be a typical 0. vulgaris. 

(14) B.M., 1924.9.9. 11-13. Octopus rugosus (Bosc), Robson (1924 and 1929). 
Natal, Gilchrist Stns. 388 and 389. C. 105-107. 

These specimens were identified but not described by Robson. The male has a 
minute ligula (Table II) and is apparently a typical vulgaris ; there is one broken 
spermatophore in Needham's organ but it was not removed. The two females have 
ripe ovaries with small eggs. The specimens are rather small in size for sexually 





Horn Coils 



. i8- 7 


. 40-50 












mature females of vulgaris but are apparently referable to this species and resemble 
the male with which they were associated. 

(15) B.M., 1926.10.20.30. Polypus granulosus (Lamarck), Massy (1927) ; Octopus 
rugosus (Bosc), Robson (1929). Cape Colony, South African Museum 1311D. 
C. 109. 

This female was described by Massy and indices based on her measurements are 
given in Table I. The specimen closely resembles the South African specimens from 
Natal, listed above, and similarly can be referred, in all probability, to 0. vulgaris. 

(16) B.M., 1889.4.24. 18. Octopus granulatus Lam., Hoyle (1886) ; Octopus rugosus 
(Bosc), Robson (1929). Challenger Coll., Simon's Bay, Cape of Good Hope, 
10-20 fms. C. 114. 

A very young specimen, with dorsal papillae in a characteristic rhomb, very 
probably referable to 0. vulgaris. 

In addition to the specimens listed above Robson's type of Octopus rugosus var. 
sanctae helenae (B.M., 1868. 3. 12. 1) was examined but nothing new can be added 
to the original description (Robson, 1929) . Although it is a badly distorted specimen 
with somewhat deceptive features there seems to be no valid reason why it should 
not be assigned to 0. vulgaris. The same is true also of Octopus verrucosus Hoyle 
(1886) (B.M., 1889.4.24. 16). The type of this species was fully redescribed by 
Robson (1929) who states that " It seems to exhibit characters of 0. vulgaris and 
0. rugosus in combination with individual peculiarities." The distinctive features 
listed by Robson do not appear to be sufficiently diagnostic to justify recognition 
of this species as distinct from 0. vulgaris. 

The following summary is offered to clarify the present synonymy of 0. vulgaris, 
in the light of the considerations outlined above. 

Octopus vulgaris Lamarck 

Octopus vulgaris, Robson (1929). All specimens from the north-eastern, north- 
western, central and south Atlantic, and from West and South Africa, which 
Robson refers to this species must be accepted as correctly identified. Speci- 
mens from the Indo-Pacific require further confirmation. 

Octopus rugosus, Robson (1929). The majority of specimens from the eastern, 
western, central and south Atlantic, and from West and South Africa which 
Robson referred to this species are probably correctly assigned to 0. vulgaris. 
It cannot be established that 0. rugosus is a valid species. Some American 
specimens are referable to 0. briar eus Robson, and 0. joubini Robson (Pickford, 
1945, 1946), and one specimen from the Cape Verde Islands, described as a new 
species in the present contribution, must be excluded. 

Octopus rugosus var. sanctae helenae, Robson (1929) = 0. vulgaris. 

Octopus verrucosus, Hoyle (1886), Robson (1929) = 0. vulgaris. 

Octopus vulgaris, Pickford (1945, 1946). The proposed synonymies of western 
Atlantic species appear to remain valid. 0. geryonea Gray, 0. eudora Gray, 


0. filosus Howell, 0. carolinensis Verrill, and various other obscure western 

forms are assimilated into the synonymy of 0. vulgaris. 

Octopus vulgaris, Adam (1952). Probably all the West African specimens which 

Adam referred to this species are correctly assigned. No doubt would arise if 

it were not for the two specimens listed from the Cape Verde Islands, but not 

identified, which must be placed in a new species (see below). 

There is an urgent need for the re-examination of specimens assigned either to 0. 

vulgaris or to 0. rugosus, granulatus or tuberculatus from the Indo-Pacific. As Robson 

has pointed out, the ligula of oriental specimens of 0. rugosus (as understood by him) 

differs from that of Atlantic specimens ; the average size is larger and the calamus 

shorter. A consideration of the data, so carefully assembled by Robson, appears to 

indicate that the Indo-Pacific rugosus-complex is composite in character and 

distinct from the Atlantic rugosus-complex. It is difficult to understand why he 

discarded this interpretation but one factor was the confusion that arose due to the 

inclusion of western Atlantic species that are not 0. vulgaris in the synonymy of 

0. rugosus. The range of variation in critical characters, such as the length of the 

ligula, was thus enlarged in such a manner as to obscure the true situation. 

Octopus vincenti n. sp. 

Octopus granulatus (part), Hoyle (1886). 
Octopus rugosus (part), Robson (1929). 
Octopus sp., Adam (1952). 

Holotype : lies du Cap- Vert, 28.x. 1948, Exped. Oceanogr. Beige cotes africaines de l'Atlantique 
Sud. Inst. Royal Sci. Naturelles de Belgique. 

Specimen examined : 

B.M., 1889.4. 24. 17. Octopus granulatus Lam., Hoyle (1886) ; Octopus 
rugosus (Bosc), Robson (1929). Challenger Coll., St. Vincent, Cape Verde 
Islands, 25. iv. 1876, depth 15-20 fms. C. 118. 

In 1952 Adam listed a male and female from the Cape Verde Islands which were 
obviously different from Octopus vulgaris, but to which he hesitated to give a new 
name. The length of the ligula in the male is quite outside the range of vulgaris 
and the horn of the spermatophore is straight, without indications of uncoiling 
(Adam, Fig. 53D). These two features alone are sufficient to justify the recognition 
of a new species since no such males have hitherto been described from the eastern 
Atlantic. In addition the animal is of small size, mantle length 25-29 mm., the arms 
are short, the web deep and the suckers small, close to the lower limits for 0. vulgaris. 
The female has a similar combination of characters but, on account of the great 
variability of Octopus vulgaris, Adam concluded that it was not possible to recognise 
the female as a distinct species. 

During the course of the present investigation it was discovered that the male 
listed above, in the Collections of the British Museum, presented the same combina- 
tion of characters. It was identified by Hoyle as 0. granulatus and referred by 
Robson to 0. rugosus, but under no circumstances can it be placed, along with other 
eastern Atlantic specimens of Robson's rugosus list, in the synonymy of 0. vulgaris. 


The size of the ligula alone precludes such an identification. The similarity to Adam's 
male is so great that there can be no doubt that the two specimens are conspecific. 
The only special feature of the British Museum specimen is the presence of a faintly 
darker area in front of the eye, about 4 mm. in diameter, which could possibly be 
interpreted as the remains of a faded ocellus ; but this interpretation seems unlikely. 

The British Museum specimen is not sexually mature and, in view of the highly 
distinctive spermatophores of Adam's specimen, which were in a good state of preser- 
vation and have been carefully figured, it seems preferable to designate this male 
as the holotype. The female, also from the Cape Verde Islands and with similar 
bodily features, is almost certainly referable to the same species but caution dictates 
that it should not be designated as the gynotype. It is unfortunate that the ovary 
was not sufficiently advanced for the size of the eggs to be determined. 

The bodily proportions and characteristics of the three specimens are summarized 
in Table III. Adam did not describe either the funnel organ or the penis. Un- 
fortunately the British Museum specimen does not allow one to give any account 

Table III. — Octopus 

vincent i n. sp. 


ular comparison of specimens. 





Holotype : $ (Adam, 1952) 
$ (Adam, 1952) 
BM.1889.4.24.17, 6" 

29 55 59 
25 72 60 
25 88 68 




34 9-5 10 6-2 40 55f 
22 9-0 10 — — — 
28 8-o 9 5 -6 40 — 

* Calculated from Adam's index (Longest arm /Mantle length). 

f Calculated from Adam's Fig. 53 D x + ^V It is not quite clear whether the two sections, horn and 
sperm reservoir respectively, show the complete spermatophore or whether a middle piece has been 
omitted. If this is the case the spermatophore length index would be greater. 

of these characters. The funnel organ is too poorly preserved for description. The 
specimen had already been dissected and no sign of the penis remains ; it was 
probably small and undeveloped since the animal is not sexually mature. There are 
no spermatophores. 

Diagnosis : A medium-sized species (mantle-length 24-29 mm.) with granular 
skin and bodily proportions similar to those of immature specimens of 0. vulgaris. 
The arms are short, 2 to 2-5 times the length of the mantle, the first arms shortest. 
The web is deep, about one-third the length of the longest arm, the D sector deepest, 
the A sector shallowest. The suckers do not exceed 10% of the mantle-length and 
there are no specially enlarged suckers in either sex. The funnel organ is not known. 
The gills have 9 or 10 primary lamellae in each demibranch ; the high number 
resembling that found in 0. vulgaris. In the male the third right arm is hectocoty- 
lized and the ligula represents 4-5-6-2% of its length, the calamus opens near the 
middle, two-fifths of the distance from the last sucker to the tip of the ligula. The 
spermatophore is about half the length of the mantle and the horn is straight, 
without coils. The size of the eggs is unknown. 

It may be enquired whether any other specimens, either among those listed by 
Adam or in the series from the British Museum can reasonably be assigned to this 
species. British Museum males have the characteristic minute ligula of 0. vulgaris. 


A juvenile male from the Cape Verde Islands (B.M., 1855.9.29.2) can also be 
excluded. The arms are longer and the web less deep, moreover the state of im- 
maturity at a mantle-length of 31 mm. indicates that it is the young of a large 
species. The immature female from the Cape Verde Islands (B.M., 1851 . 1 . 24 . 1) with 
a mantle-length of 50 mm. may be excluded for similar reasons, including also the 
larger relative size of the suckers. Among Adam's specimens there are two males 
that have an unusually large ligula for 0. vulgaris (Goree, 10. vi. 1947, LLI 3*5 ; 
Goree, 24. ix. 1946, LLI 2-9) but the spermatophore of the latter, illustrated by 
Adam (Fig. 53A), is typical of 0. vulgaris, with a spirally coiled horn. Moreover in 
both these specimens the arms are long, the web only of moderate depth, and the 
suckers large as in 0. vulgaris. Among females there is one (Goree, 31^.1950) with 
unusually short arms, a moderately deep web, and small suckers, that might well 
be referred to this species. However, the characters of the females present no clear 
cut distinctions by which one species could be separated from the other. A clarifica- 
tion of the problem must await the capture of a gravid specimen ; it may be antici- 
pated that the ripe eggs may show some distinctive features. 

It is unlikely that the sexually mature male of 0. vincenti could be confused with 
any other Atlantic species. The differences between this species and 0. vulgaris 
have already been discussed. In the western Atlantic there are four Caribbean 
species that have a ligula of approximately the same proportions as in 0. vincenti. 
0. briareus Robson is readily excluded, the arms are long, the number of gill lamellae 
low, and the spermatophore horn is coiled. 0. joubini Robson is a nearly smooth- 
skinned species with short arms, the spermatophore horn is straight but the number 
of gill lamellae is low, well below the minimum recorded for 0. vincenti. The ocellate 
species, 0. hummelincki Adam, has, in addition to this distinctive feature, a low 
number of gill lamellae and a coiled spermatophore horn. The nearest affiliation 
would appear to be with 0. burryi (Voss, 1950, 1951). This species has a characteristic 
color pattern in the form of a purplish band which extends the full length of each 
arm on the inner (dorsal) side. Apart from this highly distinctive feature, it shows 
a general resemblance to 0. vincenti and, in preparing a key for the separation of 
the two species it would be necessary to depend primarily upon the presence or 
absence of the purple arm band. Thus the arms are short, the web deep, and the 
number of gill lamellae high. The ligula is similar in size to that of 0. vincenti and 
the horn of the spermatophore is nearly straight with only a few spiral turns (one 
proximal and three distal in the one that has been described). 

The problem of 0. rugosus in the Indian Ocean is so little understood that it would 
be futile, at the present time, to offer any comparison with these or other oriental 

My best thanks are due to Dr. W. J. Rees for facilities to study the British 
Museum specimens. 


i. Twenty- two specimens from the eastern Atlantic, the west African coast and 
South Africa which Robson (1929) identified as 0. rugosus and which are now in 
the Collections of the British Museum have been re-examined. 


2. Fifteen are either juvenile or sexually immature but, with one doubtful excep- 
tion, they can reasonably be assigned to 0. vulgaris Lam. 

3. One juvenile specimen from Madeira has a peculiar zebra pattern on the arms 
and may therefore represent the young of some different species. 

4. Six of the 7 adults show no features which would justify their separation from 
0. vulgaris Lam. 

5. One adult male from the Cape Verde Islands is assigned to a new species. A 
male and female of this species were described but not named by Adam (1952) and 
Adam's male has been designated as the Holotype of Octopus vincenti n. sp. 

6. The nature of the South African vulgaris-rugosus population is discussed and it 
is concluded that, at most, the South African specimens may represent a local race 
of slightly smaller size and with other minor differences, not specifically separable 
from 0. vulgaris Lam. 


Adam, W. 1952. C6phalopodes. Exp6d. Oceanogr. Beige dans les Faux C6tieres Africaines 

de l'Atlantique Sud, Res. Sci. 3 (3) : 1-142, pis. 1-3. 
Bosc. L. 1792. Sepia rugosa. Actes Soc. d'Hist. Nat. Paris, 1 (1) : 24, PI. V, figs. 1-2. 
Hoyle, W. E. 1886. Report on the Cephalopoda collected by H.M.S. " Challenger " during 

the years 1873-76. Rpt. Sci. Res. Voyage, H.M.S. " Challenger," Zool. 16 : 1-246, pis. 1-33. 
Lamarck, J. B. 1799. Sur les genres de la Seche, du calmar et du poulpe. Mem. Soc. Hist. 

Nat. Paris, 1 : 1-25. 
Massy, A. L. 1925. On the Cephalopoda of the Natal Museum. Ann. Natal Mus. 5 : 201-230. 

1927. The Cephalopoda of the South African Museum. Ann. S. A jr. Mus. 25 : 1 51-168. 

Pickford, G. E. 1945. Le poulpe am6ricain : A study of the littoral Octopoda of the western 

Atlantic. Trans. Conn. Acad. Arts Sci. 36 : 701-81 1, pis. 1-14. 
1946. A review of the littoral Octopoda from central and western Atlantic stations in the 

collections of the British Museum. Ann. Mag. Nat. Hist. (11) 13 : 412-429. 
Robson, G. C. 1924. On the Cephalopoda obtained in South African Waters by Dr. J. D. F. 

Gilchrist in 1920-21. Proc. zool. Soc. London, 1924 : 589-686. 
1929. A Monograph of the Recent Cephalopoda. Part I — Octopodinae. 236 pp., pis. 1-7. 

London (Brit. Mus.). 
Voss, G. L. 1950. Two new species of cephalopods from the Florida Keys. Rev. Soc. Mala- 

cologica 7 (2) : 73-79, figs. 1-3. 

1951. Further description of Octopus burryi Voss with a note on its distribution. Bull. 

Marine Sci. Gulf Carib. 1 (3) : 231-240, figs. 1-6. 




Robsonella (= Joubinia) fontaniana (Orbigny) is a well defined species, endemic 
to the subantarctic or antiboreal region of South America. Supposed records of this 
species from the Indian Ocean and from the Central Pacific are subject to the gravest 
suspicion, as pointed out by Robson (1929). Massy (1925) described a specimen from 
Natal which Robson has considered to be a local variety. Two closely related but 
supposedly distinct species have been described from Australia and New Zealand, 
viz. R. australis (Hoyle) and R. huttoni Benham. Polypus campbelli of Smith is 
included in the synonymy of R. australis (Robson, 1929, footnote p. 145). 

Robson (1929) placed fontaniana and australis (+ campbelli) in the new genus 
Joubinia but this was shown to be pre-occupied and the name Robsonella was sub- 
stituted by Adam (1938). The characters of the genus were summarized by Robson 
and have been reviewed recently by Dell (1952). They are as follows : the arms are 
subequal, the web is deep and subequal, the mantle aperture is partly closed, the 
adlateral tooth of the radula is supposedly bicuspid, the penis has a long primary 
and a small secondary diverticulum, and the ligula of the hectocotylus is remarkably 
stout with inrolled sides and wide cheeks. We may disregard the characters of the 
arms and web since these are not of generic value. The partial closure of the mantle 
is also unimportant since many species of the genus Octopus have a slightly narrowed 
mantle opening of Robson's type B, moreover the mantle aperture is wide in R. 
australis (Benham, 1942). The long diverticulum of the penis is highly distinctive as 
a species character, common to all three forms, but there are species of Octopus which 
show a similar feature. The secondary diverticulum to which Robson attached great 
significance is apparently a variable feature since it was lacking in one of Dell's 
specimens of R. australis. There remains only the remarkably stout form of the ligula 
which may be conveniently retained, at least for the present, as a generic character. 
However it appears probable that the genus Robsonella is not a valid one and that 
the species which have been assigned to it should be returned to the genus Octopus. 

A second problem concerns the validity of the distinction between R. fontaniana 
and the two Australasian species. Time did not permit a re-examination of the 
specimens of R. australis (-{- campbelli) which are in the collections of the British 
Museum, but the material has been well described and we have also the accounts 
of Benham (1942) and Dell (1952) for comparison. The entire series of n specimens 
of R. fontaniana was carefully re-examined with a view to confirming and enlarging 
our concept of this South American species (Tables I and II). There are insignificant 
differences between the mean values of the indices as given by Robson (1929) and 
as determined here. In addition there are some features which call for further 



Table I. — Robsonella fontaniana (Orbigny). Bodily characteristics of eleven specimens 
in the Collections of the British Museum .* 

B.M. No. 










1851.1.24.5 . 







11 -o 



1869.6.5.54 . 










1869.6.5.65 . 









ca. 8 

1869.6.5.69 . 

6* juv. 



















1848.6. 16.2 . 










1869.6.5.63 . 










1868.7. 10.2 . 










1899. 8. 31 .84 










1869.6.5.62 . 










1851.1.24.4 . 















11 -o 



* Symbols as in. 



Table II.- 

B.M. No. 


-Robsonella fontaniana (Orbigny). Characteristics of male specimens in 
the Collections of the British Museum* 

SpLI Sp. Horn Comments 

165 No coils — 

83 One turn — 

(not formed) Immature 

(not formed) Immature 


7-8 50 

5-i 50 


3*9 56 




* Symbols as in Pickford (1945). 

Robson's sucker-diameter index does not distinguish between the diameter of the 
largest suckers of the normal series and the diameter of the specially enlarged 
suckers. It was found that such specially enlarged suckers occur only in males and 
are present in the four representatives of this sex that were examined. The penis of 
the two adult males is as figured by Robson, with an accessory knob-like diverti- 
culum. The penes in the two immature specimens are similar but they lack the 
secondary diverticula. The ligula-length index averages 6 • 5 in the two adult speci- 
mens ; as might be expected it is somewhat smaller in an immature male. 

Two spermatophores were found in the larger male (B.M., 1851.1.24.5), lodged in 
Needham's organ, and one was removed for study. It is poorly preserved and 
doubled upon itself in a knot in the middle region. The estimated length of the 
horn plus middle piece is 38 mm., the sperm reservoir measures 15 mm. The total 
length is thus half as long again as the mantle (SpLI 165). The horn is dark and 
opaque and there is no evidence of coiling. Towards its distal end, i.e. proximal to 
the knot-like tangle, the inner tube has a moniliform appearance due to a series of 
three constrictions which are probably artifacts of preservation. 

A single spermatophore was found in the smaller male (B.M., 1869.6.5.54). The 
horn plus middle piece measure 13 mm. and the sperm reservoir measures 7 mm. 


The spermatophore-length index is thus lower than in the larger specimen (SpLI 83) . 
The horn is straight except for a single widely-open turn of a spiral towards its 
distal end. 

Nearly all the females are sexually immature with small and relatively undeveloped 
ovaries. However, the reproductive ducts are well developed and have been des- 
cribed by Robson. It may therefore be inferred that the specimens are spawned 
out or in a state of sexual quiescence rather than immature. Robson calls attention 
to the distal portion of the oviduct, or vagina, which is long and stout, but I find 
that this feature is somewhat variable. One specimen (B.M., 1851.1.24.12) has a 
well developed ovary with moderately large-sized eggs, ca. 3*5 mm. in length. The 
eggs are not fully ripe and the length of the stalk could not be determined. The 
vagina of this specimen is not abnormally stout. 


Table III gives a comparison of indices and numerical characteristics of the 
species of Robsonella, compiled from data presented by Robson (1929), Benham 
(1942, 1943), and Dell (1952), and from the re-examination of the British Museum 
series of R. fontaniana. This table demonstrates the difficulty of assessing the 

Table III. — Comparison of the species that have been assigned to the genus Robsonella.* 




















R . fontaniana (Orbigny) 

B.M. Seriesf . 






11 -o 




R. australis (Hoyle) 

Robson (1929) 











" campbelli " 










Benham (1942) 










Dell (1952) 










R. huttoni (Benham) 

Dell (1952) 










* Symbols as in Pickford (1945). Doubtful specimens from the Indian Ocean and Central Pacific 

f Based on new measurements except for the arm-length index which is taken from Robson (1929) 
and which had to be used, instead of the mantle-arm index, to permit comparison with the published 
data for the other forms. 

I Specially enlarged suckers found only in males. 

|| Sexually mature specimens only. 

differences between the three species. R. huttoni differs from R. australis in its 
larger size, narrower body and much narrower head but a familiarity with the 
complexities of octopodan taxonomy suggests that these features are difficult to 
evaluate. The same is true of the relatively somewhat smaller size of the suckers. 
Both these Australasian species differ from R. fontaniana primarily in the smaller 
number of gill lamellae ; the ranges, however, overlap and the type of " campbelli " 


has fully as many lamellae as a typical specimen of R. fontaniana. It must be 
remembered that an accurate count of the number of primary lamellae is highly 
subjective, depending upon the number of minute terminal foliations that are 
included in the count. However, the difference appears to be a valid one and must 
be accepted as such in the present state of our knowledge of these species. As 
indicated by Dell (1952) R. huttoni will probably prove to fall within the range of 
R. australis, but it is also evident that the Australasian specimens are at least 
racially distinct from the South American form. 

A few more points may be discussed. The eggs of R. australis measure 2 • 5 mm. 
in length and have a short stalk (Benham, 1942). The eggs of R. huttoni are of about 
the same size, 3 mm. (Benham, 1943). Eggs, similar in size, measuring about 3 • 5 mm., 
are present in the ovary of a mature female of R. fontaniana, described here. 

The characters of the hectocotylus and penis have been fully discussed by previous 
investigators and appear to offer no evidence for separating the species. Attention 
may be called to a curious error in Dell's paper, in discussing the affinities of R. 
huttoni he states on p. 40 that " The ligula index is appreciably lower than in 
australis — average 6-7 as compared with 10 • 8." But his table (p. 34, Table 9) shows 
that his own specimens of R. australis have a low index, averaging 5*4. The specimens 
appear to have been sexually mature since the presence of a spermatophore is 
mentioned in at least one of them. 

The spermatophores of R. fontaniana are described for the first time. They are 
nearly as long as the mantle, or considerably longer. The horn is straight, as far as 
could be determined, or with a single distal turn, but admittedly the preservation 
is not good. Benham described and figured the spermatophores of R. australis. He 
states that they measured 40 mm. in length. The mantle-length of the male from 
which these spermatophores were taken is not given, but even his largest specimen 
has a mantle-length of only 36 mm. Therefore the spermatophore, like that of R. 
fontaniana, is about as long as or longer than the mantle. In the text Benham states 
that the " projectile apparatus " consists of a " closely-coiled spring". This would 
lead one to suppose that the horn of the spermatophore was coiled. However, a 
study of his figures (Benham, 1942, Figs. 6 and 7) suggests quite otherwise. The 
horn is apparently straight and what Benham mistook for the coils is clearly, in 
reality, the internal structure of the lumen which always has this delicate spiral 
structure, irrespective of whether the horn is straight or coiled (compare with the 
spermatophores of 0. macropus, figured by Pickford, 1945, PI. IV). The spermato- 
phores of R. huttoni have not been described. 

The author is indebted to Dr. W. J. Rees, for facilities to study these specimens.- 


i. Eleven specimens of Robsonella fontaniana (Orbigny) in the Collections of the 
British Museum have been redescribed. The spermatophores, described for the first 
time are nearly as long as or longer than the mantle ; the horn is straight or with a 
single spiral turn towards its distal end. The eggs are moderately large, ca. 3*5 mm. 
in length. 


2. The status of the genus Robsonella is discussed. It is considered probable that 
the two Australasian species, R. australis (Hoyle, 1885) and R. huttoni (Benham, 
1943) are synonymous but together they form an assemblage that is at least racially 
distinct from the South American species, R. fontaniana. The chief distinguishing 
characteristic is the lower average number of primary gill lamellae in the Australasian 


Adam, W. 1938. Robsonella nom. nov. fur Joubinia Robson, 1929. (Cephalopoda, Octopoda). 

Zool. Anz. 121 : 223. 
Benham, W. B. 1942. The octopodous Mollusca of New Zealand : (1) The midget Octopus 

of coastal waters. Trans. R. Soc. New Zealand, 72 : 226-236. 
1943- The octopodous Mollusca of New Zealand. — Part II. Trans. R. Soc. New Zealand, 

73 : 53-57. 
Dell, R. K. 1952. The Recent Cephalopoda of New Zealand. Dominion Mus. Bull., No. 

16 : 1-157. 
Massy, A. L. 1925. On the Cephalopoda of the Natal Museum. Ann. Natal Mus. 5 : 201-230. 
Pickford, G. E. 1945. Le poulpe americain : A study of the littoral Octopoda of the western 

Atlantic. Trans. Conn. Acad. Arts Set. 36:701-811. 
Robson, G. C. 1929. A Monograph of the Recent Cephalopoda. Part I — Octopodinae. London 

(Brit. Mus.). 


2 SEP 1955 






ZOOLOGY Vol. 3 No. 4 

LONDON: 1955 




Pp. 169-200 ; Plates 1-2 ; 41 Text-figures 

ZOOLOGY Vol. 3 No. 4 

LONDON: 1955 

(NATURAL HISTORY), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical Series. 

Parts will appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

This paper is Vol. 3, No. 4 of the Zoological series. 


Issued August, 1955 Price Ten Shillings 





Introduction ........... 171 

Classification . . . . . . . . . . -173 

Genus Epicrius Can. & Fanz. . . . . . . . -174 

Epicrius mollis (Kramer) . . . . . . . 175 

Epicrius canestrinii Haller . . . . . . . .183 

Epicrius menzeli Schweizer . . . . . . . .185 

Epicrius minor Willmann . . . . . . . .188 

Epicrius spinituberculatus n.sp. ....... 188 

Species Dubiae . . . . . . . . . .191 

Genus Berlesiana Turk . . . . . . . . . .192 

Berlesiana cirratus (Berl.) . . . . . . . -193 

Berlesiana denticulata n.sp. ........ 193 

Geographical distribution of the Epicriidae ..... 196 

Discussion ........... 196 

Summary ............ 198 

Acknowledgments .......... 198 

References ........... 198 


The family Epicriidae Berl. comprises a small group of Mesostigmatid mites readily 
distinguished in the adult stage by the polygonal network of bi- or tri-iurcate tuber- 
cles on the dorsal shield and the absence of an ambulacral apparatus on the first 
pair of ambulatory appendages. This characteristic ornamentation of the dorsum 
(" seriebus papillarum fuscarum symmetrice reticulato ") was first described as 
occurring in Gamasus reticulatus Grube, 1859, but unfortunately the original descrip- 
tion is inadequate for the recognition of its specific identity. In 1877 Canestrini 
& Fanzago figured a species for which they proposed the name Epicrius geometricus, 
thereby also erecting the new genus Epicrius. Later, Canestrini (1885) added a 
short description of the species. In the meantime, Haller (1881) had given a defini- 
tion of Epicrius and added two other species to it, Gamasus mollis Kramer, 1876 and 
Epicrius canestrinii Haller, 188 1. His description and figures of the latter are 
sufficiently detailed for recognition of the species. 

Berlese (1886-87) in "Acari, Myriopoda et Scorpiones hucusque in Italia reperta " 
re-described and figured E. geometricus stating that it was found " in totius Italiae 
muscis, praecipue in montibus." In the same work E. canestrinii is referred to as a 

ZOOL. 3, 4. 14 


variety of geometricus. Four other species, Epicrius corniger BerL, Epicrius glaber 
BerL, Epicrius laelaptoides BerL and Epicrius mollis (Kramer) Berl. were also con- 
sidered to be congeneric with E. geometricus. Later, Berlese (1916a) emended his 
concept of the genus Epicrius and transferred corniger, glaber and laelaptoides to the 
genus Lasioseius s. lat., and mollis to the genus Epicriopsis Berl. Further, in Berlese 
(1916&) Gamasus mollis Kramer 1876 non Berl., 1887 was considered to be the nymphal 
stage of E. geometricus which therefore falls into synonymy. Three new species of 
Epicrius were also described in this paper, namely, E. cirratus, E. washingtonianus 
and E. parisiensis. The latter was made the type of a new subgenus Di epicrius. 

Absolon (1899) erected the genus Eugamasus (preoccupied by Eugamasus BerL, 
1893) for Eugamasus cavernarum Absolon found on the excretory material of bats in 
a cave in Moravia. There is no doubt that this species is an Epicriid, probably of the 
genus Epicrius. It does not appear to have been found subsequently. 

George (1906) re-described three species of Epicrius from Lincolnshire. The 
species he identified as Epicrius mollis (Kramer) is Epicriopsis horridus (Kramer) 
and what he believed to be Epicrius canestrinii has recently been given the new name 
Cornubia georgei by Turk (1943) . The specimens of the third species are in the collec- 
tions of the British Museum (Natural History) and are correctly determined as 
Epicrius geometricus Can. & Fanz. (= E. mollis). 

Schweizer (1922) listed two species of the genus Epicrius from Switzerland, 
E. geometricus and Epicrius menzeli Schweizer, 1922. The specimens of the former 
were collected from three main regions, namely, " Mittelland," " Jura " and" Alpen". 
The females described from " Mittelland " and " Alpen " were correctly determined, 
but the female described and figured from the " Jura " was, as will be shown below, 
Epicrius canestrinii Haller. 

Tragardh (1942) in his revision of the genus Epicrius unfortunately deals only 
with E. mollis. This paper is important for the historical account of the genus and 
the description of the larva and the protonymph of the genotype. In addition, 
Tragardh has dealt with the classification of the family Epicriidae in several papers 
on the comparative morphology of the Mesostigmata e.g. Tragardh 1938, 1946a & b. 

Turk (1943) has also added to our knowledge of the Epicriidae in his work on the 
British species. He erected a new genus, Berlesiana, for Epicrius cirratus Berl. on 
the basis of the structure of the sternal region in the female. The genus Cornubia, 
which was also proposed in this paper, was later made a synonym of Ameroseius BerL 
(Turk, 1953). Two species are included in the genus Epicrius, namely, E. mollis 
and E. geometricus. His E. mollis, however, is Epicriopsis horridus (Kramer) and 
this invalidates the discussion in that paper on the synonymy of E. mollis and E. 

The most recent work on the genus Epicrius is by Willmann (1953). A new 
sub-genus, Epicriella, was proposed for Epicrius [Epicriella) minor Willm. 

The object of the present work is to review our knowledge of the classification of 
the Epicriidae and to re-describe and figure, when possible, the known species. 
This revision is based on material in the collections of the British Museum (Natural 
History), the Oudemans Collection at Leiden and Dr. Jos. Schweizer 's Collection at 


Basle. Dr. G. Lombardini has also kindly undertaken comparisons of specimens of 
two species with the types in the Berlese Collection at Florence. 


The supra-generic classification used in this paper follows that given by Evans 
(1955) in which the Epicriina was divided into two super-families, the Epicrioidea 
and Zerconoidea. 


" Mites with the epigynal portion of the genital plate in the female reduced to a 
narrow chitinized rim overlapping the genital orifice. Male genital opening situated 
in the sternal shield in the region of coxae II and III and closed by two plates, the 
anterior of which bears a pair of hairs. Chelicerae dentate in both sexes but without 
spermatophoral process in the male. Pedipalps with five free segments, specialized 
seta on palptarsus two or three pronged" (Evans, 1955). 

Super-family Epicrioidea Evans 

Epicrioidea Evans, G. O. Bull. Brit. Mus. (nat. Hist.) 1954, Zool. 2 : 295. 

Dorsal shield in both sexes provided with bi- or trifurcate tubercles forming a 
polygonal network, and a pair of large dorso-lateral protuberances of unknown 
function. Dorsal setae simple, setose or strongly barbed. Both sexes with jugularia; 
sternal setae II and /or IV may be situated on a sclerotised shield or on the inter- 
scutal membrane. Geniti-ventral shield in the female large, flask-shaped or rect- 
angular. Sterniti-genital shield in the male extending posterior to coxae IV. 
Gnathosoma with four pairs of ventral setae ; corniculi short, digitiform. Pedipalps 
with the tibia and tarsus fused dorsally. Specialized seta on palp tarsus three- 
pronged. Peritreme markedly reduced or absent. Stigmata enclosed in the lateral 
extension of the dorsal shield. Leg I without ambulacral apparatus. Tarsus I (and 
sometimes tibia I) with specialized, clubbed sensory setae. Legs II-IV with multi- 
lobed pulvillus and two claws. 

Family Epicriidae Berl. 

Epicriidae Berlese, A. Bull. Soc. ent. Ital. 1885, 17 : 129. 

The only family of the Epicroidea and therefore with the above characters. The 
Epicriidae consists of two genera which may be distinguished as follows: 

I. The majority of the dorsal setae more than ioo\l in length, simple or setose. 

Dorso-lateral protuberances large, conspicuous. Tarsus I with three or 

more clubbed sensory setae ...... Epicrius Berl. 

-. The majority of the dorsal setae less than 60 \l in length, stout and strongly 

barbed (fig. 32). Dorso-lateral protuberance small, inconspicuous. 

Tarsus I with less than three pairs of clubbed sensory setae . Berlesiana Turk 


I. Genus Epicrius Canestrini & Fanzago 

Epicrius Canestrini, G. & Fanzago, F., Atti. 1st. Venet. 1877, (5) 4 pt. 1 : 131. 

Eugamasus Absolon, Ph. C. K., Zool. Anz. 1899, 22 : 324. 

Parasejus Tragardh, T. Naturw. Untersuch. des. Sarekgebirges. 1910, 4, 4 : 432. 

Epicrius (Diepicrius) Berlese, A., Redia, 1916, 12 : 151. 

Epicrius (Epicriella) Willmann, C, Sitzber. osterr. Akad. Wiss. math-nat. Kl. 1953, 1, 6 : 474. 

Dorsal setae simple or setose ; the majority longer than 100/1. Dorso-lateral 
protuberance large. Geniti-ventral shield flask-shaped. Tarsus I with three or 
more clubbed setae. Other characters as in the definition of the super-family. 

Type : Epicrius geometricus Can. & Fanz., 1877, (= Gamasus mollis Kramer, 1876). 

This genus contains five species which may be considered valid and three species of 
uncertain status pending the re-examination of the type material. The former may 
be separated according to the following key to both sexes : 

Key to the species of the genus Epicrius Can. & Fanz., 1877 


1. Sternal setae II situated on the interscutal membrane between the jugularia and a 

shield bearing sternal setae III and IV .... Epicrius minor Willmann 
-. Sternal setae II situated on a well-sclerotised shield with setae III or setae III and 

IV 2. 

2. Dorsal setae D2 and D4 about one-third the length of setae D3 . . . . 3. 
-. Dorsal setae D2 to D4 approximately equal in length. Sternal setae II to IV on an 

undivided shield. Interscutal membrane between geniti-ventral and anal shields 

with seven setae ........ Epicrius menzeli Schweizer 

3. Projections of dorsal tubercles with rounded extremities (PI. 1) .... 4. 
-. Projections of dorsal tubercles sharply pointed at their extremities (PI. 2). Sternal 

setae II to IV on an undivided shield. Interscutal membrane between geniti- 
ventral and anal shields with four setae. Anal shield with the usual three setae 

Epicrius spinituberculatus sp. n. 

4. The seta posterior to the large dorso-lateral protuberance separated from it by a 

transverse row of tubercles. Sternal setae II to IV on an undivided shield. The 
four setae between the genitiventral and anal shields each situated on a platelet 1 

Epicrius canestrinii Haller 
-. The seta posterior to the large dorso-lateral protuberance not separated from it by a 
row of tubercles. Sternal setae IV (metasternals) separated from shield bearing 
setae II and III. The pair of setae between the geniti-ventral and anal shields not 
on platelets ........ Epicrius mollis (Kramer) Berl. 


1. Anal shield completely fused with the dorsal shield. Posterior margin of the 
sterniti-genital shield strongly convex. With three pairs of setae on the interscutal 
membrane posterior to the sterniti-genital shield . . Epicrius mollis (Kramer) Berl. 

-. Anal shield free, or partly fused with the dorsal shield. Posterior margin of the 

sterniti-genital shield truncated . . . . . . . . . 2. 

1 In some specimens the platelets on one side of the mid-ventral line may be fused so that there are 
only three platelets between the geniti-ventral and anal shields, (see p. 13). 


2. Setae D2 and D4 about one-third the length of setae D3 . . . . . 3. 
-. Setae D2 to D4 approximately equal in length. Ventri-anal shield with ten pre-anal 

setae .......... Epicrius menzeli Schweizer 

3. Projections of dorsal tubercles rounded at their extremities. Region between the 

sterniti-genital and anal shields occupied by a large rectangular shield bearing 
three pairs of setae ....... Epicrius canestrinii Haller 

-. Projections of dorsal tubercles sharply pointed. With a ventri-anal bearing three 

pairs of pre-anal setae ...... Epicrius spinituberculatus sp. n. 

Epicrius mollis (Kramer), 1876. 

Gamasus mollis Kramer, P. Arch. Naturgesch. 1876, 42 : 82 (fig.) ; Berlese, A., Redia, 1916, 
12 : 150. Epicrius mollis, Tragardh, I., Ark. Zool. 1942, 34a (4) : 3 (fig.). 

Epicrius geometricus Canestrini, G. & Fanzago, F. Atti 1st. Venet. 1877, 1 : 60 ; Haller, G. 
Arch. Naturgesch. 1881, 47 : 190 ; Canestrini, G. Prosp. Acarof. Ital. 1877, (5) 4 : 131 
(fig.) ; Berlese, A., A. M.S., Padova, 1886, 30 : 8 (fig.) ; George, C. F., Naturalist, 1906 : 265 

The writer follows Berlese (1916) and Tragardh (1942) in considering G. mollis 
to be the nymphal (deutonymphal!) stage of E. geometricus. The two main objec- 
tions to the acceptance of this synonymy are that Kramer's description and figure 
are not sufficiently detailed for the specific identity of the species and that the 
original figure of mollis shows only one pair of dorso-lateral protuberances whereas 
geometricus is figured with two pairs. It is probable, however, that Berlese examined 
the type material of geometricus before coming to any decision regarding the syno- 
nymy of the species so it is assumed that the original figure of geometricus is 
inaccurate. Berlese (1886) figured geometricus with one pair of dorso-lateral pro- 
tuberances and certainly no Epicriid with two pairs of these structures has been 
described since Canestrini & Fanzago (1887). 

The following re-description of E. mollis is based on material collected over a period 
of twelve months from litter under a Spruce stand in South Bedfordshire. 1 

Dorsal shield. The dorsal surface of the larva is incompletely covered by a weakly 
sclerotized shield bearing nine pairs of setae distributed as in Text-fig. 1. These 
setae are probably homologous with those occuring on the anterior shield of the larvae 
of the free-living Laelaptoidea (Evans, 1953). The interscutal membrane posterior 
to the shield is granulated and bears three pairs of long whip-like setae. In the pro- 
tonymph the dorsum is completely covered by a weakly sclerotized shield (Text-fig. 2) . 
The chief features of interest in the chaetotaxy of the shield are : 

(1) the addition of five pairs of dorsal setae (D) making a total of ten pairs ; and 

(2) the addition of setae ni in the anterior half of the shield. 

The relative lengths of the setae Di to D5 are approximately the same in both 
stages, i.e. D2 and D4 are considerably shorter than Di, and D3. Setae D6 and D7 
in the protonymph are much shorter than D7, D9 and Dio. The writer has been 
unable to distinguish, with certainty, between larval and protonymphal setae in the 
posterior half of the shield. The sclerotized dorsum of the deutonymph is charac- 
terized by the development of a pair of large dorso-lateral protuberances (Text-fig. 3). 

1 Collected during investigations financed by H. M. Forestry Commission whilst the writer was on the 
staff of Rothamsted Experimental Station, Harpenden, Herts. 





The relative lengths of the setae Di to Dio are approximately the same as in the 
protonymph. The deutonymphal setae in the anterior half of the shield are indicated 
by the symbol n2. There is no increase in the number of setae on the dorsal shield 

Text-figs. 4, 6. Epicrius mollis (Kr.). Fig. 4, dorsum of female. Fig. 6, venter of 

female. Abbreviations as Fig. 1. 

from deutonymph to adult (Text-fig. 4). The dorsum of the adult (both sexes) is 
strongly sclerotized and characteristically ornamented. The ornamentation com- 
prises bi- and trifurcate tubercles forming a polygonal network (PI. 1). The dorsal 
setae are setose (Text-fig. 5). The chaetotactic pattern is shown in the figure. 
Setae D8 which are considerably shorter than Do, in the deutonymph are almost 

ZOOL. 3, 4. 



equal in length to them in the adult. The inornate region surrounding the dorso- 
lateral protuberance carries two setae and a large pore-like structure. 

Ventral shields. The ventral surface of the larva is weakly sclerotized. The 
sternal shield, extending from the posterior margin of coxae I to the posterior margin 
of coxae III bears three pairs of simple setae. The oval anal shield is provided 
with a pair of para-anals and a post-anal seta. These setae are long and extend 
beyond the posterior margin of the body. The interscutal membrane posterior to 
coxae IV has four pairs of setae of which two pairs are situated between the sternal and 
anal shields. The sternal and anal region in the protonymph is similar to that in the 
preceding stage. The region between the sternal and anal shields is provided with 
three pairs of setae. The sternal shield in the deutonymph bears four pairs of setae 
and is more strongly sclerotized than in the larva and protonymph. The anal 
shield has three long setae. 

The venter of the female is shown in Text-fig. 6. The first pair of sternal setae are 
situated on small platelets (jugularia) surrounded by striated membrane. Sternal 
setae II and III stand on a distinct shield situated between coxae II and III and the 
metasternals on separate shields postero-lateral to it. The separation of setae IV 
from the main sternal shield is difficult to detect in specimens not cleared in lactic 
acid. The sternal shield is not fused with the endopodals. The geniti- ventral shield 
is large and flask-shaped, and bears two pairs of setae. The epigynal portion of the 
shield is poorly developed being reduced to a narrow semi-transparent band along 
the anterior margin of the shield. The region between the geniti-ventral and the 
ventri-anal shields has a pair of simple setae. The ventri-anal bears a pair of pre- 
anals and para-anals, and a post-anal seta. The para-anals and the post-anal seta 
are situated behind the anal opening. There is a pair of conspicuous pore-like 
structures on the lateral margin of the shield. The remaining sclerotized structures 
of the venter are a bi-furcate tritosternum, a porose plate situated posterior to coxae 
IV and the ventro-lateral extension of the dorsal shield, which is provided with a 
network of tubercles and a number of setae. The interscutal membrane in the region 
of the jugularia and lateral to the coxae is richly provided with short spines. 

The vental surface of the male differs considerably from that in the female (Text- 
fig. 7). The jugularia are present in normal position, but the region between coxae II 
and IV is occupied by a well-sclerotized shield bearing four pairs of setae and three pairs 
of " pores/' This shield, the sterniti-geniti- ventral, extends beyond the posterior 
margin of coxae IV. It is truncated anteriorly but strongly convex posteriorly. 
It is not fused with the endopodals. The male genital orifice is situated between 
coxae III and is covered by two plates. A pair of setae protrude from under the 
postero-lateral margin of the anterior plate. The ventri-anal shield is fused with 
dorsal shield, but the setae associated with the anal region are present in the same 
position as in the female. The interscutal membrane posterior to the sterniti- 
geniti- ventral shield carries three pairs of setae. 

Gnathosoma, pedipalps and chelicerae. The ventral surface of the gnathosoma 
in the larva is provided with two pairs of setae, namely, the rostrals and the external 
posterior rostrals (Text-fig. 8). The corniculi are slender and digitiform. There are 



four rows of denticles in the region of the ventral groove. The pedipalps comprise 
five free segments. The palptibia and tarsus are divided by a distinct suture ven- 
trally, but are fused dorsally. The specialized seta on the palptarsus is three- 
pronged. The palptrochanter has a spur-like projection ventro-laterally. The 
chaetotactic formula for the trochanter, femur and genu is (0-4-5). The ventral 

Text-figs. 5, 7, 12, 13. Epicvius mollis (Kr.). Fig. 5, distal end of dorsal seta. Fig. 7, 
venter of male. Fig. 12, palptibia and tarsus (ventral) of female. Fig. 13, chelicera of 

surface of the gnathosoma in the protonymph bears two additional pairs of setae — 
the internal posterior rostrals and the capitular setae (Text-fig. 9). The chaeto- 
tactic formula for the palptrochanter, femur and genu is (1-4-5) . In the deutonymph 
there is a further increase in the number of setae on the pedipalp. These additional 
setae are indicated by the symbol n2 in Text-fig. 10. There is no change in the 
chaetotaxy of the pedipalp from deutonymph to adult (Text. fig. n). The structure 


of the venter of the palptibia and tarsus is shown in Text-fig. 12. These segments 
are fused dorsally. The development of the chaetotaxy of the gnathosoma and first 

Text-figs. 8, 9, 10, 11. Epicrius mollis (Kr.). Gnathosoma (ventral) of larva (Fig. 8), 
protonymph (Fig. 9), deutonymph (Fig. 10) and female (Fig. 11). v., rostral setae ; 
ext.p.r., external posterior rostral setae ; int. p. v., internal posterior rostral setae ; 
cap., capitular setae. 

three free segments of the pedipalp in Epicrius is essentially similar to that described 
by Evans (1953) for the genus Typhlodromus Scheuten {Gamasina — Laelaptoidea) . 

The tectum in all stages is broadly triangular in shape and has a denticulate 
margin (Text-figs. 8-1 1). There appears to be considerable variation in the degree of 
denticulation, at least in the series examined. 



The chelicerae are chelate-dentate. 

structure of the chelicera of the female is shown in Text-fig 

The dentition is weak in all stages. The 


Stigmata and peritr ernes. The stigmata and peritremes are not developed in the 
larva. In the protonymph the stigma (st) is situated ventro-laterally in the region 
of the third intercoxal space (Text-fig. 14). The peritreme (p) is markedly reduced 
and is enclosed with the stigma in an oval depression of the body. The main tracheal 
branch is conspicuous. The stigma and peritreme is the same in the deutonymph. 

Text-figs. 14, 15. Epicrius mollis (Kr.). Fig. 14, stigmal region of protonymph. 
Fig. 15, stigmal region of female, st., stigma ; p., peritreme ; tr., trachea. 

In the adult, however, the stigma is enclosed in the heavily sclerotized lateral exten- 
sion of the dorsal shield. The stigmal opening is situated on a small protuberence 
(Text-fig. 15). The only indication of a peritrematal plate (?) is the presence of a 
well-defined line running from the stigma to the level of coxa I and thereby separating 
a strip of smooth sclerotized plating from the ornamented dorsal shield. 

Legs. All the legs comprise six free segments with the terminal segment incom- 
pletely separated into a metatarsus and tarsus. Leg I is the longest in all stages 
and the tarsus is without an ambulacral apparatus. In the adult, tibia and tarsus I 
are provided with specialized sensory (?) setae ventro-laterally (Text-fig. 16). These 
long setae, of which there is one on the tibia and four on the tarsus, are club-like 
distally. The swollen head of the seta is minutely setose. The number and position 
of these setae are different in the species examined, but since the majority of the 
specimens were poorly preserved — the distal end of the setae are easily broken off 
in prepared specimens — full use could not be made of this character. The remaining 



ambulatory appendages terminate in a multi-lobed pulvillus and two claws (Text- 
fig. 17). The majority of the setae on the femur to tibia are stout and spinose, and 
stand on distinct bases. 

~ ^L^L/^ _ 


Text-figs. 16, 17. Epicrius mollis (Kr.). Fig. 16, Tibia and tarsus I. Fig. 17, 

Ambulacral apparatus of Leg. II. 
Text-fig. 37. Berlesiana denticulata sp. n. Tarsus I. 


Larva, 265-280/4 in length and 1 95-200/4 in breadth. 
Protonymph, 310-340/4 in length and 245-250/6 in breadth. 
Deutonymph, 440-490/4 in length and 305-335/^ in breadth. 
Female, 640-660/4 in length and 405-41 o/£ in breadth. 
Male, 570-600/4 in length and 360-370/4 in breadth. 


Locality. This species is the most abundant and widely distributed of the genus. 
It has been recorded from moss and forest litter in Italy, Austria, Switzerland, 
Germany, Holland, Sweden and the British Isles. 

Epicrius canestrinii Haller, 1881 

Epicrius canestrinii Haller, G., Arch. Naturgesch. 1881, 47 : 191 (fig.) ; Oudemans, A. C, 

Zool. Anz. 1939, 126 : 307. 
Epicrius (Diepicrius) parisiensis Berlese, A., Redia, 1916, 1 12 : 152, syn. nov. 
Epicrius geometricus, Schweizer, J. Verh. Natf. Ges. Basel, 1922, 33 : 46 $. 

The following re-description of E. canestrinii is based on specimens of both sexes 
in the British Museum (Nat. Hist). — from the Michael Collection. The specimens 
are mounted in Canada Balsam. 

Female. The dorsal shield, about 500/^ in length and 350/4 in breadth, shows 
the normal network of tubercles. The latter are bi- or trispinate ; the projections 
having rounded extremities. The dorsal chaetotaxy is shown in Text fig. 18. 
Setae D2 and D4 are considerably shorter than setae D3, but setae D7, D8 and Do, 
are approximately equal in length. The dorso-lateral protuberance is well-developed 
and broadly triangular in shape. The ornamentation in the region of the protuber- 
ance differs from that in the other species of the genus examined by the writer. The 
anterior tubercle seta and the dorso-lateral protuberance are distinctly separated 
from the posterior tubercle seta by a transverse row of tubercles. The anterior 
seta is about twice the length of the posterior seta. 

Ventrally, the tritosternum and jugularia are normal for the genus (Text-fig. 19). 
Sternal setae II, III and IV are situated on a rectangular shield extending from the 
middle of coxae II to the middle of coxae III. The anterior margin of this shield, 
in the majority of the specimens examined, is concave and the posterior margin 
convex. The geniti- ventral shield is of the usual form with two pairs of simple setae. 
The epigynal portion of the shield is well-developed. The interscutal membrane 
between the geniti- ventral and anal shields bears four pairs of setae. The four setae 
lying between the geniti- ventral and anal shields each lie on a platelet. The number 
of platelets appears to be variable. In the female figured by Schweizer (1922) and 
which the writer has examined, the two platelets on one side are fused. The anal 
shield is small and provided with three setae, of which the para-anals are situated 
behind the anal opening. 

The gnathosama and pedipalps are normal for the genus. A critical examination 
of the chelicerae could not be carried out owing to the state of preservation of the 

The legs are also normal for the genus. It was not possible to be certain of the 
number of clubbed setae on the terminal segments of the first pair of legs. As far 
as could be seen three clubbed were present on the tarsus. 

Male. The ornamentation and chaetotaxy of the dorsal shield is similar to that 

1 Volume 12 of Redia was published on 191 7, but separates of Berlese's paper containing the 
description of this species are stated to have been published on 23.viii.1916 (see p. 177). 



in the female. The chief differences between the sexes is to be seen in the structure 
of the ventral surface. 

The elongate sterniti-genital shield, posterior to the paired jugularia, has four pairs 
of setae and two pairs of " pores." It extends from the middle of coxae II to beyond 
the posterior margin of coxae IV. The posterior margin of the shield is truncated 

O 100 200/U 


Text-figs. 18, 19, 20. Epicrius canestrinii Haller. Fig. 18, dorsum of female. Fig. 
19, venter of female. Fig. 20, venter of male,, anterior tubercle seta ; 
post, tub.s., posterior tubercle seta. 

(Text-fig. 20). The region between the sterniti-genital and anal shields is occupied 
by large rectangular shield bearing three pairs of setae. Either side of this shield, 
on the interscutal membrane, are two setae. The anal shield, partly fused with 
the dorsal shield, is about as broad as long and has three setae and a pair of pore- 
like structures. The para-anals lie on the level with the posterior margin of the 
anal opening. 

Dimensions. Female : 520-550/* in length, 340-352/* in breadth. Male; 
480-520/4 in length, 297-320/* in breadth. 



Locality. The type locality is in the vicinity of Bern, Switzerland. It occurs in 
moss and decaying vegetable material and has subsequently been recorded from the 
following areas : Meudon (nr. Paris), France (Berlese, 1916) ; Dissenhofen and 
Jouxtal, Switzerland (Schweizer, 1922) ; Porth Gwarra, Cornwall, England (Michael 
Coll.) ; and Delden, Holland and Sucy-en-Brie, France (Oudemans Coll.). 

This species may be readily separated from other species of the genus by the struc- 
ture and chaetotaxy of the region of the dorso-lateral protuberance. Haller (1881), 
in his original description of the species, deals only with the structure of the dorsum. 
He gives a figure of dorso-lateral protuberance and associated setae showing the 
separation of the posterior tubercle seta from the tubercle (Text-fig. 21). This 

O^ <&<&<& of- 


st.tub.s. <&o 



^oj post.tub.s. 

Text-figs. 21, 22. Epicrius canestrinii Haller. Ornamentation and chaetotaxy in 
the region of the dorso-lateral protuberance of the female. Fig. 21, after Haller 
(1881). Fig. 22, specimen in the Michael Collection. Abbreviations as in Fig. 18. 

figure agrees very well with the one based on material examined by the writer (Text, 
fig. 22). 

The male of Epicrius (Diepicrius) parisiensis Berlese agrees in all details with the 
male of E. canestrinii. 

Epicrius menzeli Schweizer, 1922. 
Epicrius menzeli Schweizer, J. Verh. Natf. Ges. Basel, 1922, 33 : 47, figs. $ & $. 

The following re-description of E. menzeli is based on the type material kindly 
lent to the writer by Dr. J. Schweizer, Birsfelden. 

Female. The dorsal shield, 748/6 x 45 1#, is provided with a network of tubercles 
as in other species of the genus. The tubercles are bi- and trispinate. The setae 
of row D are considerably more uniform in length than in the preceding species — 
setae D2 and D4 being approximately equal in length to D3. The dorso-lateral 
protuberance is strongly formed and the setae lying immediately anterior and 
posterior to it are contained in the same area. 

Ventrally, the tritosternum is normal for the genus (Text-fig. 23). The first pair 
of sternal setae are situated on distinct platelets. The remainder of the sternal 
setae lie on a large sternal shield extending from the middle of coxae II to the middle 



of coxae III. Sternal setae II are widely separated from setae III and IV, which lie 
in a transverse line along the posterior margin of the shield. A pair of distinct 
" pores " lies between setae III and IV. The geniti- ventral shield is large and 
flask-shaped, with two pairs of setae. The epigynal portion of the shield extends a 
considerable distance beyond the genital orifice. The geniti-ventral is faintly 
sculptured and has a pair of " pores." The interscutal membrane between the geniti- 
ventral and the anal shields bears seven setae arranged as in the figure. The anal 




Text-figs. 23, 24. Epicrius menzeli Schweizer, female. Fig. 24, gnathosoma 

(ventral). Fig. 23, venter. 

shield has three setae and a pair of large pore-like structures (muscle attachments (?)). 
The para-anal setae are situated posterior to the anal opening. The metapodalia 
are small, as are the porose-plates posterior to coxae IV. 

The gnathosoma and pedipalps are normal for the genus (Text-fig. 24). The 
rostrals (240/4), the interior posterior rostrals (7Jfi), the exterior posterior rostrals 
(66/i) and the capitular setae (no/^) are distributed as in the figure. The chelicerae 
are chelate-dentate and normal for the genus. The legs conform in general with those 
in the preceding species. Tibia I (approximately 185/4 in length) is provided with 
one clubbed seta and tarsus I (approximately 220/1) with three clubbed setae. 

Male, The dorsal shield in this sex measures 627-638/4 in length and 418-429/4 



in breadth. The ornamentation and chaetotaxy is basically the same as in the 
female (Text-fig. 25). 

Ventrally, the jugularia are well-developed and widely separated from the sterniti- 
genital shield (Text-fig. 26) . The latter has four pairs of simple setae and two pairs 
of " pores " distributed as in the figure. The shield is incised anteriorly and truncate 

Text-figs. 25, 26. Epicrius menzeli Schweizer, male. Fig. 25, dorsum. Fig. 26, venter. 

posteriorly. It is faintly sculptured. The genital orifice is situated between coxae 
III. The genital sclerites are as in other species of the genus. The region posterior 
to the sterniti-genital is occupied by a large ventri-anal shield bearing thirteen simple 
setae and a pair of conspicuous " pores ". The para-anal setae lie posterior to the 
anal opening. The shield is faintly sculptured. The porose platelets are situated 
postero-lateral to coxae IV. The metapodalia are apparently absent. 


The gnathosoma and pedipalps are essentially the same as in the female. The 
chelicerae were not fully visible in the preparations examined. 

Tibia I (176-181/4 in length) is provided with one clubbed seta and tarsus I (178- 
198/4 in length) with three clubbed setae. 

Dimensions. Female: 748/4 in length, 451/4 in breadth. Male : 627-638/4 in 
length, 418-429/^ in breadth. 

Locality. The type material comprises one female and three males collected 
in damp Beech leaves in a ditch near Bennwil (" Basler Jura "), Switzerland. 

Epicrius minor Willmann, 1953 

Epicrius (Epicriella) minor Willmann, C, Sitzber. osterr. Akad. Wiss. math-nat. Kl. 1953, 
1, 6 : 474, fig. ?. 

This species is characterized by the structure of the sternal region of the female, 
the only sex known. Sternal setae II lie on the interscutal membrane between the 
jugularia and an elongate shield bearing setae III and IV. The geniti- ventral shield 
is large and flask-shaped with two pairs of setae and a pair of " pores ". The inter- 
scutal membrane between the geniti-ventral and the anal shields has one pair of 
setae only. The anal shield bears five setae, of which the paired para-anals are 
situated behind the anal opening. The details of the ornamentation and chaetotaxy 
of the dorsal shield are not given in the original description of the species. 

Dimensions. Female 405/4 in length, 255/4 in breadth. 

Locality. " Gipel des Unterberges, etwa 1,300 m., in den Voralpen von Nieder- 
osterreich, Waldquadrat in niederem Buchenkrummholz mit zwischenstehenden 
jungen Fichten . . . Untervegetation bedeckt den Boden zu einem Drittel, viel 
Buchenfallaub., 1 $ " (Willmann, 1953). 

Epicrius spinituberculatus sp. n. 

Female. The dorsal shield, 594/4 in length and 396/^ in breadth, is richly pro- 
vided with tubercles forming a distinct network. The structure of the tubercles is 
unique amongst the genus Epicrius in that the distal projections are sharply pointed 
and not obtuse as in the other species (Text-fig. 27 and PL II). The number of 
dorsal setae is normal for the genus. Setae D2 and D4 are considerably shorter 
than D3, whilst D7, D8 and D9 are approximately equal in length (Text-fig. 28). 
The dorso-lateral protuberance is well-developed and the setae lying anterior and 
posterior to it are contained in the same area. 

Ventrally, the tritosternum is normal for the genus (Text-fig. 29). The first pair 
of sternal setae are situated on distinct platelets, whilst sternal setae II to IV are on a 
sclerotized shield extending from the middle of coxae II to the middle of coxae III. 
The distribution of setae II to IV is similar to that in E. menzeli. The geniti-ventral 
shield is flask-shaped, being constricted in the region of the anterior of two pairs of 
setae which are situated on it. The epigynal portion of the shield is similar to that in 
E. mollis. The striated interscutal membrane between the geniti-ventral and anal 
shields is provided with two pairs of simple setae. The anal shield is not fused 



with the dorsal shield and bears three setae. The para-anals are situated behind 
the anal opening. A pair of large pore-like structures is present antero-laterally on 
the anal shield. The metapodalia and porose plates are present in the positions 
shown in the figure. 





Text-figs. 27, 28, 29. Epicrius spinituberculatus sp. n., female. Fig. 27, dorsal tubercles. 
Fig. 28, dorsal shield. Fig. 29, venter 



The gnathosoma and pedipalps are normal for the genus. The chelate-dentate 
chelicerae were not sufficiently exposed for critical study. 

The general structure of the legs is normal for the genus. Tibia I, 187/6 in length, 
has one clubbed seta. The number of clubbed setae on tarsus I, which is approxi- 
mately equal in length to tibia I, could not be ascertained owing to the state of pre- 
servation of the specimen. 





Text-Fig. 30. Epicrius spinituberculatus sp. n. Venter of male. 

Male. The network of tubercles and the relative length of setae D2-D4 on the 
dorsum of the male are the same as in the female. The lateral and postero-lateral 
setae appear to be relatively longer in length, but this may be due to the distortion 
of the dorsal setae in the female as the result of mounting. 

Ventrally, the tritosternum and jugularia are normal for the genus. The sterniti- 
genital shield bears four pairs of simple setae and two pairs of " pores " (Text- 
fig. 30). The genital orifice is situated between coxae III. The region posterior 
to coxae IV is occupied by a large ventri-anal shield similar in shape to that in 
E. menzeli, but having only three pairs of pre-anal setae. The para-anals are 


situated posterior to the anal opening. A pair of large pore-like structures is 
also present on the shield. The interscutal membrane in the region of the coxae 
is richly provided with minute spines. 

The gnathosoma and pedipalps are normal for the genus. The chelicerae are 

Tibia I, 154/4 in length, is provided with one clubbed seta and tarsus I, 165/4 in 
length, has three of the four macro-setae clubbed. 

Dimensions : Female : 597/4 in length, 396/4 in breadth. Male : 535-540/4 in 
length, 340/4 in breadth. 

Locality : A single female (holotype, 1930. 8. 25. 2198) labelled Epicrius canes- 
trinii Haller in the Michael Collection in the British Museum (Nat. Hist.) The spec- 
imen is without a precise locality, as are the majority of Michaels' specimens. 
Also two males (Allotype, 1954.9.8.2 and Paratype 1954.9.8.3), collected by the 
writer from a thick layer of humus (F-layer) under bracken in the Leri Valley, near 
Dol-y-bont, Cardiganshire, Wales, on 19. iv. 1954. 

E. spinituberculatus may be readily separated from other species of the genus by 
the structure of the dorsal tubercles, the chaetotactic pattern of the dorsal shield, 
and the chaetotaxy of interscutal membrane between the geniti-ventral and anal 
shields in the female. 

Species Dubiae 

The following three species which probably belong to the genus Epicrius, have not 
been examined by the writer ; the descriptions are inadequate for their certain 

Gamasus reticulatus Grube, 1859 

Gamasus reticulatus, Grube, A. E., Arch. Naturk. Liv. Ehst.-u. Kurl. 1895, (2) 1 : 459 and 

Epicrius reticulatus, Oudemans, A. C, Zool. Anz. 1939, 126 : 306. 

The original description is sufficient only to indicate that this species probably 
belongs to the genus Epicrius. The ornamentation of the dorsal shield of the unique 
specimen is described as " seriebus papillarum fucarum symmetrice reticulato ". The 
first pair of legs are without claws. The type locality is in Latvia. Grube's material 
may be in Dorpat or Breslau (Oudemans, 1939). 

Eugamasus cavemarum Absolon, 1899 

Eugamasus cavemarum, Absolon, Ph. C. K., Zool. Anz. 1899, 22 : 324. 
Epicrius cavemarum, Oudemans, A. C, Zool. Anz. 1939, 126 : 307. 

The description of this species is chiefly based on the male. The ventral surface 
in this sex is said to be divided into a number of plates. The sternal plate is small 
(or narrow) and rounded off, whilst the genital plate, connected with the anal plate 
behind, is prolonged and rounded off anteriorly. Further the " abdominalplatte " 
is divided into two distinct plates. The remainder of the description deals with 


structures which are characteristic of all other species of the genus. The description 
of the female is too short to be of value in defining the species. 

If Absolon's description of the ventral surface of the male is correct, then E. 
cavernarum is undoubtedly a valid species. The writer is inclined, however, to treat 
it as a species dubia until the type material can be re-examined. 

E. cavernarum was found on the excreta of bats in a cave (Slouperhole) in Moravia, 

Epicrius washingtonianus Berlese, 1916 

Epicvius washingtonianus Berlese, A., Redia. 1916, 12 : 151. 

Berlese opens his description of this species by stating that " species hanc describere 
bene non possum . . . ". The description deals with the ornamentation and chaeto- 
taxy of the dorsal shield only and is as follows : 

". . . Tamen a caeteris hucusque notis est diversum, quod corniculis dorso- 
lateralibus (ad quartos pedes) caret et areolis minus numerosis quam in E. cirrato 
gaudet. Pili dorsi (qui adhuc persistunt) breves, nulla barbula ornati, simplices, 
forsitan omnes intersese statura subaequales (ad 8o/£ long.). Tuberculi dorsi 
plerumque bilobi, rarius trilobi. Ad 6oo/£ long. ; 420/^. lat. Caeteris hucusque 
notis specibus maior. 

Habitat. Inveni in muscis ad ' Washington ' collectis." 

II Genus Berlesiana Turk, 1943 

Berlesiana Turk, F. A., Ann. Mag. nat. Hist. 1943, (11), 10 : 855. 

This genus was proposed by Turk for Epicrius cirratus Berlese. He considered the 
structure of the sternal region in that species to be so distinctive that " when the 
types are re-examined a new family will have to be made ". This would appear to 
be likely in the light of Tragardh's concept of the family Epicriidae (Tragardh, 1939). 
The discovery of a new species undoubtedly congeneric with E. cirratus but with 
sternal setae II and III in the female on an undivided shield shows that the structure 
of the sternum in the Epicriidae is more variable than Tragardh realised, and 
cannot be used as a major character in the classification of the Epicriina. 

The ornamentation of the dorsal shield and the structure of the dorsal setae in 
E. cirratus separates it from the mollis — group of species. The writer proposes to 
retain Berlesiana Turk as a valid genus but at the same time emending the original 
definition as follows : 

With the general characters of the genus Epicrius but differing from it in 
the following characters : tubercles of the dorsum more numerous ; postero- 
lateral protuberance poorly developed ; dorsal setae short, stout and strongly 
barbed ; geniti- ventral shield in the female more or less parallel-sided ; tarsus 
I with less than three pairs of clubbed setae. 
Type : Epicrius cirratus Berl., 1916. 


The genus Berlesiana contains two species which may be separated by the structure 
of the sternal shield in female, as follows : 

1. Sternal setae II to IV each situated on a separate shield Berlesiana cirrata (Berl.) 
-. Sternal setae II and III on an undivided shield . . Berlesiana denticalata sp. n. 

Berlesiana cirrata (Berl.), 1916 

Epicrius cirratus Berlese, A., Redia. 1916, 12 : 151. 

Berlesiana cirratus Turk, F. A., Ann. Mag. nat. Hist. 1943, (n) 10 : 855. 

In the original description of this species Berlese mentions the ornamentation 
and chaetotaxy of the dorsal shield. The dorsal setae are about 50/* in length. 
The dorso-lateral protuberance is stated to be absent, but it is possible that Berlese 
may have overlooked it, since a poorly developed protuberance is present in the 
normal position in the species described below. 

The sternal plate in the female is described as "in partes duas laterales fracto ; 
quaeque pars autem fissura transversa plus minusve bene in scutulis duobus rotundis, 
in medio piliferis divisa ". The ventri-anal shield is provided with five setae in the 
female. The male is not described. 

Dimensions. Female : 440/* in length, 300/4 in breadth. Male : " considerably 
smaller ". 

Locality. This species is known from the type localities only ; the female from 
Chianti and Pontedera, and the male from near Genoa. In all localities the species 
was found in moss. 

Berlesiana denticulata sp. n. 

Female. The dorsal shield, 415-425/* in length and 260-265/t in breadth, 
completely covers the dorsum of the mite. The ornamentation of the shield con- 
sists of numerous tubercles, predominantly tri-lobed (Text-fig. 31). The dorso- 
lateral protuberance is not nearly as well developed as in the genus Epicrius. The 
dorsal setae with the exception of four pairs of simple setae antero-laterally, are 
short (44-55/*), stout and strongly barbed (Text-fig. 32). Setae Di to Dio are 
approximately equal in length. The distribution of " pores " is shown in the figure. 

Ventrally, the tritosternum and jugularia are as in the genus Epicrius. Sternal 
setae II and III are situated on an undivided shield extending from the middle of 
coxae II to the middle of coxae III (Text-fig. 33). Setae IV (the metasternals) lie off 
this shield. In the holotype the metasternal seta on one side lies on the inter-scutal 
membrane but on the other side there are two setae situated on a small shield. This is 
undoubtedly an aberration since in the paratypes sternal setae IV are situated on the 
interscutal membrane postero-lateral to the shield bearing setae II and III. The 
geniti-ventral shield is ornamented with punctations and is roughly rectangular in 
shape. Its posterior margin is truncated. The epigynal portion of the shield is 
well-developed and denticulate along its anterior margin. The geniti-ventral bears 
two pairs of simple setae. The endo-podal plates are poorly developed and not fused 
with the sternal shield. The ventri-anal shield is elongate, truncate anteriorly and 



provided with a pair of pre-anal setae. The surface of the shield is scabrid. There 
is a tendency for increased setation of the ventri-anal. In one of the paratypes 
there are four pre-anal setae. The ventri-anal is not fused with the dorsal shield 



^&^S @D 

« d^Sfc^ @D3 

s Oi,<?1)Q % ©D5 

<b o J 
^> ©D9 


Text-figs. 31, 33. Berlesiana denticulata n. sp., female. Fig. 31, dorsal shield. 

Fig. 33, venter. 

(Text-fig. 34) . A pair of simple setae is present between the geniti- ventral and anal 

The gnathosoma is provided with four pairs of setae ventrally of which the 
rostrals are considerably longer than the other three pairs. The pedipalps have 



five free segments. The chaetotaxy of the palptrochanter, femur and genu is 
(2-5-6) . The palptarsus is incompletely separated from the tibia ; the two 
segments being fused dorsally. The specialized seta on the palptarsus is three- 

Text-figs. 32, 34, 35, 36, 38. Berlesiana denticulata n. sp. Fig. 32, dorsal seta of 
female. Fig. 34, postero-ventral region of female. Fig. 35, tectum of female. Fig. 36, 
chelicera of female. Fig. 38, venter of male. 

pronged. The tectum is denticulate and basically the same as in species of 
Epicrius (Text-fig. 35). The chelicerae are chelate-dentate (Text-fig. 36). 

Leg I is considerably longer than the body and is without an ambulacral apparatus. 
Tibia I, about no/i in length, has no clubbed setae. The chaetotaxy of tarsus I is 
shown in Text-fig. 37 (p. 182). This segment has two clubbed setae and, proximally 


a long stout seta lying parallel to the longitudinal axis of the segment. Legs II to IV 
terminate in a pulvilms and two claws, and are of the form in the genus Epicrius. 

Male. The ornamentation and chaetotaxy of the dorsal shield in the male is 
essentially similar to that of the female. 

Ventrally, the jugularia are relatively large and are distinct from the sterniti- 
genital shield which extends from the middle of coxae II to well beyond the posterior 
margin of coxae IV. This compound shield bears four pairs of simple setae distri- 
buted as in Text-fig. 38. The genital orifice is situated between coxae III and its 
structure is normal for the family. The surface of the shield appears to be corrugated 
in preserved specimens. It is not strongly sclerotized. The ventri-anal shield is 
similar in form to that in the female except that it bears an additional pair of pre-anal 
setae. The striated interscutal membrane between the sterniti-genital has a pair 
of simple setae. There is also evidence of increased setation of the ventral shields 
in the male. In one specimen two short setae were present posterior to the last 
pair of " sternal " setae on the sterniti-genital. 

The gnathosoma, pedipalps and legs are basically the same as in the female. 

Dimensions. Female : 415-425^ in length, 260-265^ in breadth. Male : 400- 
405/^ in length, 240-250/^ in breadth. 

Locality. Five females and seven males from litter under Beech at Boxhill, 
Surrey (collected by Messrs. S. K. Eltringham and K. H. Hyatt, 21.viii.1953). 
Holotype female, 1954 . 9 . 8 . 9 ; Allotype male, 1954. 9. 8. 10; Paratypes, 1954.9.8. 


Any discussion on the zoogeography of the free-living Acarina is open to criticism 
on the basis that very little work has been done on them outside the Palaearctic and 
Nearctic Regions. It is, therefore, not possible to be dogmatic about the limits of 
distribution of any group of free-living mites. Bearing this in mind, it can be stated 
that, at present, the Epicriidae are recorded only from the Palaearctic and Nearctic 
Regions ; ten " species " from the former and only one species from the latter. 1 Epi- 
crius mollis (Kr.) appears to be the most widely distributed species in the Palaearctic 
Region having been recorded from a number of localities between Sicily in the south 
and Sweden in the north. 

The writer has been unable to find representatives of this family in recent collec- 
tions of free-living mites from Uganda, India, Singapore and Rennell Is. 


The examination of the above mentioned species of the genera Epicrius and Berle- 
siana enables one to discuss in some detail the value of the morphological characters 
which have formed the bases of the various classifications of the Epicriidae put 
toward by previous workers. In several of these investigations, particularly those 

1 Baker & Wharton (1952) state that " on Guam Epicrius sp. was found among the rhizomes of epi- 
phitic ferns ..." in which case this is the first record of the family outside the Palaearctic and Nearctic 


of Tragardh, on the comparative morphology of the Mesostigmata, the classification 
has been built up on the evidence supplied by the critical examination of only one 
species, Epicrius mollis (Kr.). The following outline of the development of the 
classification of the family since its erection by Berlese (1885) will serve as the basis 
for discussion. 

Berlese (1885) considered the genus Epicrius and Podocinium Berl. to constitute a 
a subfamily (!) Epicriidae of the family Gamasidae. The characteristic features of 
this subfamily were the elongate first pair of ambulatory appendages and the orna- 
mentation of the body of the mites. Epicrius and Podocinium were separated on the 
degree of development of the peritreme. Later (Berlese, 1892), Epicrius was placed 
with Zercon Koch and Seiodes Berl. in the Zerconidae, apparently on account of the 
position of the genital orifice in the male. This classification was retained by Berlese 
(1913) in his introduction to the " Acarotheca Italica," except that the Zerconidae 
was divided into the Zerconini and the Epicriini. Vitzthum (1929, 1931 and 1941) 
followed, in the main, Berlese's classification and at the same time emphasized the 
close relationship between the Epicriidae and the Zerconidae. 

Tragardh (1938), in the first of his important contributions to the classification 
of the Mesostigmata, separated the Epicriidae and the Zerconidae ; placing the former 
in the Sejina (= Liroaspina) and the latter in the Gamasina on differences in the 
structure of the genital and sternal regions in the females. The Epicriidae was 
regarded as a primitive group because of the absence of a distinct epigynal shield and 
the segmentation of the sternal region. The Sejidae ( = Liroaspidae) and Epicriidae 
were separated as follows : 

"A. Sternal shields III and IV fused, forming a narrow transverse shield, 

separated from the remaining sternal shield .... Fam. Sejidae. 

A A. All sternal shields fused ...... Fam. Epicriidae." 

In 1946 (19460:) the same general classification of the Liroaspina was given except 
that the sternal region of the Epicriidae was now described as " sternal shields I free, 
shields II-IV fused." But after examining the structure of the genital region in 
the male, the Epicriidae was removed from the Liroaspina to the Epicriina (Tra- 
gardh, 1946&). At the same time the Epicriina was stated to resemble the Zer- 
conina in this character. Therefore, the final result of Tragardh's researches were 
to separate the Epicriidae and the Zerconidae into distinct cohorts, chiefly on the struc- 
ture of the genital region in the females. 

The writer disagreed with Tragardh's interpretation of the structure of the genital 
region in the female of the Epicriidae and could find no evidence for his statement 
that an epigynal shield is not developed in this group (Evans, 1955). The geniti- 
ventral shield in both Epicrius and Berlesiana are basically the same as that in the 
Laelaptoidea. This is shown in Text-figs. 39-41, where a comparison is made between 
the genital region of two species of Laelaptoid mites and E. mollis. The writer 
therefore, suggested the inclusion of the Epicriidae and Zerconidae in the Epicriina, 
following Vitzthum (1941). 

The second important morphological character used by Tragardh in the classifi- 
cation of the Liroaspina and Epicriina is the structure of the sternal shield in the 


females. It is on the basis of the degree of fragmentation of the sternal shield in 
the female that the Epicriidae was originally separated from the Liroaspidae and, 
later, the Liroaspidae from the Epicrosejidae (Tragardh, 1952). This separation of 
the families appears to have been made after the examination of one species from each 
family. From the present revision of the Epicriidae it is quite obvious that the frag- 
mentation of the sternal region in this group shows considerable variety and can no 

•P- 39 

Text-figs. 39-41. Lateral view of the sterniti-geniti- ventral region of the females of 
two species of Laelaptoid mites (Figs. 39 and 40) and Epicrias mollis (Kr.), (Fig. 
41). ep., epigynal portion of the geniti-ventral shield ; g-v-sh., geniti-ventral shield. 

longer be used as a major character in the classification of the family. The following 
types of sternal " shields " were encountered in the females : 

1. Sternal setae II free ; III and IV on a shield, (E. minor). 

2. Sternal setae II and III on a shield ; IV free, (E. mollis, B. denticulata). 

3. Sternal setae II, III and IV on a shield, (E. canestrinii, E. menzeli.. 
E. spinituberculatus) . 

4. Sternal setae II, III and IV each on separate shield, (B. cirrata). 
Jugularia are present in both sexes in all species. 


The generic concept in the Epicriidae also has the structure of the sternal region 
in the females as its basis. In the recent classification given by Baker & Wharton 
(1952) the genera Epicrius and Berlesiana are placed in separate families ; the former 
in the Epicriidae and the latter in the Liroaspidae. In the present classification 
both genera have been included in the Epicriidae and the subgenera Diepicrius 
and Epicriella have been relegated to the synonymy. 


1. A revision is given of the family Epicriidae with a discussion on the classifica- 
tion of the Epicrhna. 

2. Two genera, Epicrius and Berlesiana, are considered valid. The former 
contains five valid species and three of uncertain status, and the latter two species. 

3. Keys to species are given for the identification of both sexes of Epicrius and the 
females only of Berlesiana. Epicrius {Diepicrius) parisiensis Berl. is considered to be 
a synonym of Epicrius canestrini Haller. 

4. The following two species are described as new : Epicrius spinituberculatus 
and Berlesiana denticulata. 


The writer is extremely grateful to Dr. Jos. Schweizer for the loan of material 
from his collection, to Dr. L. van der Hammen for specimens from the Oudemans 
Collection, to Dr. G. Lombardini for comparing specimens with those in the Berlese 
Collection, and especially to Dr. H. W. Parker for his criticisms of the manuscript. 

The photographs are the work of Mr. M. G. Sawyers. 


Absolon, Ph. C. K. (1899). Uber die fauna der Hohlen des mahrischen Devonkalkes (Vor- 

laufige Mittheilung) . Zool. Ariz. 22, 324. 
Baker, E. W. & Wharton, G. W. (1952). An Introduction to Acarology. Macmillan. 

465 pp. 
Berlese, A. (1882-1892). Acari, Myriopoda et Scorpiones hucusque in Italia reperta. Padova. 

(1885). Acarorum Systematis. Bull. Soc. ent. Ital. 17, 129. 

(1913)- Acarotheca Italica, Systema — Acarorum Genera. Firenze. 

(1916a). Centuria Prima di Acari Nuovi. Redia, 12, 32. 

(19166). Centuria Seconda di Acari Nuovi. Redia, 12, 150. 

Canestrini, G. & Fanzago, F. (1886). Intorno agli Acari Italiani. Atti. 1st. Venet. (5) 4, 


— ■ — (1885). Prospetto dell' Acarofauna Italiana. Padova, 60. 
Evans, G. O. (1953) • Some mites of the genus Typhlodromus Scheuten, 1857 from S.E. 
Asia. Ann. Mag. nat. Hist. (12), 7, 615. 

( I 955). A collection of Mesostigmatid mites from Alaska. Bull. Brit. Mus. {nat. Hist.) 

Zool. 2, 9 ; 287-307. 

George, C. F. (1916). Lincolnshire Mites, Epicrius. Naturalist, 265. 

Grube, A. E. (1859). Verzeichniss der Arachnoiden Liv-, Kur- und Ehstlands. Arch. 

Naturk. Liv.-Ehst.-u. Kurl. 1, 459 and 474. 
Haller, G. (1881). Acarinologisches. Arch. Naturgesch. 47, 190. 
Kramer, P. (1876). Zur Naturgeschichte einiger Gattungen aus Familie der Gamasiden. 

Arch. Naturgesch. 42, 82. 


Oudemans, A. C. (1939). Neue Funde auf dem Gebiete der Systematik und der Nomenklatur 

der Acari V. Zool. Anz. 126, 306. 
Schweizer. Jos. (1922). Beitrag zur Kenntnis der terrestrischen Milbenfauna der Schweiz. 

Vevhandl. der Naturf. Ges. Basel, 33, 47. 
Tragardh, I. (1910). Acariden aus dem Sarekgebirge. Naturw. Under such, des Sarekge- 

birges, 4, 432. 

(1938). Further contributions towards the comparative morphology and classification 

of the Mesostigmata. Ent. Tidskr. 59, 123-158. 

(1942). Zur Kenntnis der Gattung Epicrius Berlese. Ark. Zool. 34a, 4, 3. 

(1946a). Outlines of a new classification of the Mesostigmata based on comparative 

morphological data. Lunds Univ. Arsskr. 42, 1-37. 

(19466). Contributions towards the comparative morphology of the Mesostigmata 

(Acarina) VII. Ent. Tidskr. 67, 88-108. 

(1952). Acarina collected by the Mangarevan expedition to South Eastern Polynesia . . . 

(Mesostigmata). Ark. Zool. (2) 4, 46-90. 

Turk, F. A. (1943). Studies of Acari. — I. The British Species of Epicriidac, with Descriptions 

of a new Genus and two new Species. Ann. Mag. nat. Hist. (11), 10, 855-860. 
Vitzthum, H. G. (1929). Die Tierwelt Mitteleuropas, VII Acari. Leipsig. 

(1931)- Kiikenthals Handbuch der Zoologie, 3, 2, 143. 

(1941)- Acarina in Bronn's Tierreich 5, Abt. 4, Buch 5, 778. 

Willmann, C. (1953). Neue Milben aus den ostlichen Alpen. Sitzber. dsterr. Akad Wiss. 
math.-nat., Kl., 1, 6, 474. 

Ornamentation of the dorsal shield in Epicrius mollis (Kramer), x 1000. 

Ornamentation of the dorsal shield in Epicrius spinituberculatus sp. n. x 800. 


2 SEP 1955 



2 SEP 1955 



1 uah 1956 THE MONK SEALS 

(Genus Monachus) 




ZOOLOGY Vol. 3 No. 5 

LONDON : 1956 




Pp. 201-256 ; Pis. 3-8 ; 12 Text-figures 

ZOOLOGY Vol. 3 No. 5 

LONDON: 1956 

(NATURAL HISTORY), instituted in 1949, is 
issued in five series, corresponding to the Departments 
of the Museum, and an Historical series. 

Parts will appear at irregular intervals as they 
become ready. Volumes will contain about three or 
four hundred pages, and will not necessarily be 
completed within one calendar year. 

This paper is Vol. 3 , No. 5 of the Zoological series. 


Issued January, 1956 Price Eighteen Shillings 

THE MONK SEALS (Genus Monachus) 



Synopsis ............ 204 

Synonymy ............ 204 

M . monachus ........... 205 

History ........... 205 

Distribution ........... 206 

Taxonomy ........... 209 

Description . . . . . . . . . . . 210 

M. tropicalis ........... 215 

History . . . . . . . . . . .215 

Distribution . . . . . . . . . . .216 

Taxonomy ........... 218 

Description . . . . . . . . . . . 218 

M. schauinslandi ........... 220 

History ........... 220 

Distribution ........... 222 

Description ........... 223 

Specimens Examined .......... 223 

Osteology ....... .... 226 

Comparison of skulls . . . . . . . . .226 

Dentition ........... 232 

Supernumerary bones . . .. . . . . .236 

Scapula . . . . . . . . . . . 237 

Humerus ........... 239 

Radius and ulna .......... 239 

Manus ............ 239 

Pelvis and sacrum . . . . . . . . 239 

Femur . . . . . . . . . . .241 

Patella . . . . . . . . . . .241 

Tibia and fibula . . . . . . . . . .241 

Pes . . . . . . . . . . . .241 

Vertebral column .......... 242 

Ribs ............ 245 

Sternum . . . . . . . . . . 245 

Growth ............ 246 

Relationships . . . . . . . . . . .251 

Acknowledgments .......... 252 

Bibliography ........... 252 

zool. 3, 5. 




An account is given of the history, distribution, features and habits of the three species of 
monk seal Monachus monachus, M. tropicalis and M. schauinslandi. 

A list is given of the skulls, skeletons, skins and stuffed animals that have been examined. 

The skull, dentition and skeleton are described and compared with those of other Phocidae. 
The members of the genus Monachus are shown to be much more nearly allied to the Southern 
than the Northern Phocids. 

The relative growth rates of the various parts of the skull are given, and show that the facial 
region grows more quickly than the cranial region. 

The genus Monachus — the monk seals — consists of only three species. M. monachus 
(Hermann 1779) lives in the Mediterranean area, M. tropicalis (Gray 1850) in the 
West Indies, and M. schauinslandi Matschie 1905 in the Hawaiian Islands. They 
are interesting animals, but because of their widely scattered distribution, 
remoteness and scarcity, not a great deal is known about them. In this paper an 
attempt will be made to bring together as much information about the monk seals 
as it has been possible to find. 


Genus MONACHUS Fleming, 1822 

Monachus Fleming, 1822, Philos. Zool. 2 : 187 (footnote). Phoca monachus Hermann. 
Pelagios F. Cuvier, 1824, Mem. Mus. H.N. Paris, 11 : 196. Phoca monachus Hermann. 
Pelagius F. Cuvier, 1826, Diet. Set. Nat. Paris, 39 : 550. For Pelagios. 
Pelagus McMurtrie, 1834, Cuvier's Animal Kingdom, 71. For Pelagios. 
Pelagias Gray, 1837, Mag. Nat. Hist., N.s. 1 : 582. For Pelagios. 

Pelagocyon Gloger, 1841, Gemeinn. Naturgesch. 1, 34 : 163. Phoca monachus Hermann. 
Rigoon Gistel, 1848, Nat. Thier, fur hohere Schulen. X. New name for Pelagios F. Cuvier. 
Heliophoca Gray, 1854, Ann. Mag. Nat. Hist. 13 : 201. Heliophoca atlantica Gray = Phoca 
monachus Hermann. 

I. Monachus monachus (Hermann, 1779) 

Phoca monachus Hermann, 1779, Besch&f. Berlin Ges. Naturf. Freunde, 4 : 501, pis. 12, 13. 

Phoca albiventer Boddaert, 1785, Elench. Anim. 1 : 170. 

Phoca bicolor Shaw, 1800, General Zoology. 1,2: 254. 

Phoca leucogaster Peron & Lesuer, 18 16, Voy. aux Terres Austr. 2 : 47 (footnote). 

Phoca hermanni Lesson, 1828, Diet. Class. Hist. Nat. 13 : 416. 

Monachus mediterraneus Nilsson, 1838, K. Svenska Vet. Ah. Handl. 1837 : 238. 

Phoca crinita Menis, 1848, II Mare Adriatico, 153. 

Heliophoca atlantica Gray, 1854, Ann. Mag. Nat. Hist. 13 : 202. 

II. Monachus tropicalis (Gray, 1850) 

Phoca tropicalis Gray, 1850, Cat. Mamm. in colln. Brit. Mus. part 2. Seals, 28. 
wilkianus Gosse, 1851, A Naturalist's Sojourn in Jamaica, 308 (footnote). 

III. Monachus schauinslandi Matschie, 1905 

Monachus schauinslandi Matschie, 1905, Sitzungs-Berichte der Gesellschaft Nat. Freunde, Berlin, 


I. Monachus monachus 


The monk seal appears to have been always well known to the inhabitants of the 
Mediterranean area. Clark (1952) notes that bones of this seal were found in upper 
palaeolithic levels at Grimaldi, and there is evidence that seal hunting was quite an 
important activity in classical Greece. The skins were collected partly for the 
superstitions surrounding them and partly for use as clothes by the poor fisher folk. 
Boots, tents and dresses made of skin were said to protect one from lightning ; a 
seal skin drawn round a field and then hung up by the door would save the field from 
hailstones, and the right flipper was supposed to be a cure for insomnia when put 
under the head at night. Greek writers such as Plutarch, Pliny, Homer and 
Aristotle knew and wrote of the seal. Aristotle must have examined one with care 
as he gives quite an accurate account in the Historia Animalium, but Keller (1887) 
says that on the whole the poets found the animal horrible and deformed, and 
objected to its oily smell. The presence of seals gave rise to many stories. The 
" half animal fisheaters " on the west coast of Africa were said to have made a pact 
with the seals not to interfere with each other's fishing, and in the Odyssey a woman 
who died on board ship was thrown overboard to serve as food for the seals and 
fishes. Because of their love of sun and sea the seals were put under the protection 
of Phoebus Apollo, the sun god, and Poseidon, the sea god. The seal was frequently 
shown alive in these times. Even then their docility and intelligence were noted, 
and Pliny tells how he saw some which answered by growls when their names were 
called. In earlier days there must have been more monk seals round the Greek 
coasts than there are now, as several towns have taken the name of the seal. Phocis 
is the name of an ancient Greek district that stretched past Mount Parnassus to the 
Gulf of Corinth, and there is at the present time Foca at the north end of the Izmir 
Gulf, Turkey, and Foca, an inland town in Yugoslavia. Keller (1887) notes that 
very ancient pre-Darian coins (ca. 500 B.C.) show a picture of a seal, and that coins 
from Rhodes show a seal's head. He also mentions that etymologically the Greek 
work for " phoca " means the swollen or plump animal, and must have been first 
applied, though not in a taxonomic sense, to the monk seal. 

Some of the later history has been summarized by Monod (1932) . An Italian map 
of the fourteenth century shows an island between Lanzarote and Fuerteventura in 
the Canary Islands called Ya de Uegi marini — the island of sea wolves — to-day 
called more simply Lobos Island ; and in 1341 Niccoloso di Recco included seals in 
an inventory of the Canary Islands. In the fifteenth century Portuguese explorers 
found numerous bands of monk seals and killed many for their skins, and in 1434 
Alfonso Baldaya started the industrial exploitation of the seals in the bay Rio del 
Oro, north of Cap Blanc. A traveller named Zarco reached Madeira in 1418 and 
named a small cove Camara de Lobos because of the seals he found there. 
Rondoletius published his Aquatilium Historia in 1554 and included a drawing and a 
short Latin description of " de Vitulo maris mediterranei ", but the first specific name 
was given by Hermann (1779) who described an animal from the Dalmatian coast 


and called it Phoca monachus. Buffon (1782) mentions that seals from the Adriatic 
were kept in captivity in France and Germany in 1760, and describes a female seal 
that was on exhibition in Nimes in 1777. He also gives a detailed description of a 
male seal that was caught on 28th October, 1777, " off the island of Guarnero " in 
the Adriatic. He saw this seal in Paris in December, 1778, and from the similarity 
of his description of it and of its locality and habits it seems that it must have been 
the same animal described by Hermann — a fact which Cuvier noted as early as 1813. 
Hermann saw the seal in Strasbourg at the end of October and beginning of 
November, probably in 1778, and says that it was being taken on tour by a company 
of Venetians who were going to Paris to show it to the King. He mentions that the 
seal was said to have been captured in the autumn of 1777 in the Dalmatian Sea at 
Ossero. Ossero is on the island of Cherso in the Gulf of Quarnero, off the Dalmatian 
coast. Buffon notes that the seal died in August, 1779. 

Since that time there have been many references, though mostly of the occurrence 
of single individuals. Cuvier (1813) describes a seal that was kept in captivity for 
two years in a very small bath, only a foot longer and two feet wider than the animal 
itself. In this remarkably small enclosure it spent 9-10 hours of each day in six 
inches of water that were drained off each night. The London Zoological Gardens 
has three times kept a monk seal for short periods. The first specimen, a young 
female, arrived in May, 1882, but died the same day ; the second was in 1894, when 
a young animal from Madeira lived there for three and a half months ; and the last 
was in 1910, when a second animal from Madeira lived for four months. In 1926 
the American Museum of Natural History received skins and skeletons of three seals 
from the Desertas Islands ; these were believed to be the first Mediterranean seals 
in American museums. More recent references have again been concerned mainly 
with isolated occurrences, and there is at the time of writing a male monk seal from 
near Oran that has been in captivity for about two years in the Jardin des Plantes 
in Paris. 

Distribution and abundance 

The Mediterranean monk seal is known from the shores and islands of the 
Mediterranean and the western coast of North Africa (figs. 1 and 2). It has been 
recorded from : Gulf of Almeria, Spain ; Cabrera, Balearic Is. ; Toulon ; Corsica ; 
Cape Teulada and the Gulf of Cagliari in Sardinia ; the island of Pelagosa in the 
Adriatic ; the Gulf of Quarnero and Fort Opus in Yugoslavia along the Dalmatian 
coast ; the Gulf of Salonika in north Greece ; Cape Caliacra, and generally in the 
Black Sea and Bosphorus ; Tantoura ; El Arish ; Port Said ; the island of Galite 
off the Tunisian coast ; Oran ; Madeira and the Desert a Grande Islands ; the 
Canary Islands, and along the African coast, including Cap Barbas, Baie d'Etoile 
and Baie du Levrier, to Cap Blanc. 

Budker (1945) says that the southern limit along the African coast is 20 49' N. 
(approximately the latitude of Cap Blanc), that its limit of distribution is influenced 
by the temperature of the sea and corresponds with the 20 C. winter isotherm. In 
the British Museum collections there is an Ascarid from a monk seal taken in Senegal 




(approximate latitude 15 N.). It has been in the Museum since 1863, but there is 
no other information about the host. 

Apart from the few references given by Monod (1932) for the fourteenth and 
fifteenth centuries there do not seem to be any other accounts of large scale 
commercial exploitation of the monk seal, although during a visit to Madeira in 

Fig. 2. — Map of Mauritanean coast and Atlantic islands, and inset of Cap Blanc to show 

distribution of M. monachus. 

1945, Cadogan (1945) heard that " in 1943-4 a consignment of some twenty-five 
seal pelts had been seized by the Customs Officials at Funchal, and it was not thought 
likely that any further attempt would be made to commercialize them for the 
present ". Any commercial use of an animal necessarily implies its abundance in 
the area. Admiral W. H. Smyth, writing of the period 1810-1824 (in Flower, 1932) 
says that " between Alexandria and Benghazi , . , we found fish and seals in 


abundance ", but from then up to the present day there have been references only 
to single animals or small groups. Barcelo (1875) said that seals were very common 
on the shores of the Balearic Islands at the time he wrote, but Cabrera writing in 
1914 notes their disappearance from these parts. Aharoni (1930) notes that the 
seal is quite often offered for sale by fishermen from Askalon and Jaffa. Bertram 
(1943) says that a seal was seen off El Arish in about 1941, and according to Monod 
(1945), Agacino (1950) and Postel (1950) there must still be reasonably large colonies 
along the western coast of Africa down to Cap Blanc. Sixty seals were seen by 
Postel just north of Cap Blanc, and twenty-one by Agacino along the coast of Rio de 
Oro, and Cousteau and Dumas (1953) note the presence of a herd of about two 
hundred seals at Port Etienne. An adult female was surprised and killed in a cave 
in Corsica in 1947 (Troitzky, 1953), and Butler (in litt.) saw a seal in the Gulf of 
Salonika in 1950. There is still a small breeding colony in the more isolated regions 
of the Desertas Islands, although the fishermen there regard it as an enemy to their 
livelihood. A recent report in The Times (nth June, 1954) says that the monk seal 
still appears yearly on the Turkish coast. The main stronghold of the monk seal at 
present seems to be along the coast of Rio de Oro, and if not molested it is probable 
that its numbers will be maintained there. 


The first specific description of the monk seal is that given by Hermann in 1779. 
He named as Phoca monachus a male animal captured at Ossero, on the island of 
Cherso, off the Dalmatian coast in 1777, and seen by him in Strasbourg. He gives 
a very detailed description of the seal and its habits, and also a drawing of the 
whole animal with details of the head and flippers (PL 4, a). In 1782 Buffon, not 
knowing of Hermann's work, described the same seal which was, in 1778, on show 
in Paris. He gives a very good drawing of the animal (PL 4, b) but only refers to it 
as " Le phoque a ventre blanc ". These two descriptions form the basis of 
practically all the succeeding names which have been applied to the monk seal. 

Boddaert, in 1785, used Buffon's paper as the basis for his very brief Latin 
description of Phoca albiv enter, a specific name which was in use for many years. 
In volume two of the third edition of Pennant's Quadrupeds, published in 1793, 
Pennant gives a description of the Pied Seal, mentioning Buffon, but adding " This 
I saw at Chester ; it was taken near that city in May, 1766 ". It is difficult to 
know whether Pennant was applying Buffon's description to a seal actually taken 
near Chester, Cheshire, in which case it was very unlikely to have been a monk seal, 
or whether he was confusing the name with Cherso on the island of that name in the 
Adriatic. The drawing he gives is not very good, (PL 4, c) and shows an animal 
with a broad white ring round its neck and a white spot behind one flipper, the rest 
of the body being black. Pennant also describes the Mediterranean seal, quoting 
Hermann, but giving no picture. Shaw in 1800 uses Pennant's Pied Seal and 
Buffon's Phoque a ventre blanc for his Phoca bicolor, and used Phoca monachus for 
the Mediterranean seal. He uses the drawings previously given by Buffon and 
Pennant and labels both of them " Pied Seal var," 


In a brief footnote, in which he quotes a passage from Buff on, Peron (1816) uses 
the name Phoca leucogaster, and in another equally brief footnote Fleming (1822) 
first suggests the use of the generic name Monachus, " Some seals, as Ph. monachus, 
are said to have four incisors in each jaw. Such will probably be constituted into a 
new genus, under the title Monachus ". In 1824 Cuvier suggested Pelagios as a 
new generic name for Phoca monachus Hermann, a name which has given rise to 
many variants ; Cuvier himself in 1826 called it Pelagius, McMurtrie (1834) used 
Pelagus, and Gray (1837) use d Pelagias. Lesson in 1828 renamed Hermann's 
species Phoca hermanni, and Nilsson in 1838 called it Monachus mediterraneus. In 
1 841 Gloger called the monk seal of the Mediterranean Pelagocyon monachus, and in 
1848 Menis used the name Phoca crinita for an animal presumably from the Adriatic 
(book not seen). Gistel, also in 1848, proposed the generic name Rigoon instead of 
Pelagios of Cuvier. Giebel (1848) linked the monk seals with the southern Phocids 
under the genus Leptonyx and the subgenus Leptorhynchus . This part of his 
classification is given here in full : 

" Leptonyx 

a. Stenorhynchus 

1. L. serridens 

2. L. leopardinus 

b. Leptorhynchus 

3. L. weddellii 

4. L. rossii 

5. L. monachus." 

The last synonym was given by Gray in 1854, when he named the new genus and 
species Heliophoca atlantica on the skin and skull of a young animal from Deserta 
Grande, Madeira. The skull and stuffed skin of this animal are now in the British 
Museum collections (Reg. No. 1853. 10.6.4, 1063a.) 


Fully grown adults are about 8-9 ft. long. Gavard (1927) gives the length of a 
female which had produced a pup as 2-42 m. (7-9 ft.), but does not say whether 
this length included the hind flippers or not ; Troitzky (1953) gives the nose to tail 
length of an old female as 278 m. (9*1 ft.), and both Postel (1950) and Agacino 
(1950) note that the biggest animals that they saw were about 3 m. (9*8 ft.) long. 
Monod (1945) gives the length of a male animal as 2*9 m. (9-5 ft.). Of the two 
adult skins in the Museum collection the length of the male is 2*38 m. (7.8 ft.) and 
of the female 2 • 1 m. (6 -9 ft.). These measurements are from nose to tip of tail, but 
should be regarded with caution as that of the male was taken from a rather crumpled 
dressed skin, and that of the female from an undressed but folded skin. The weight 
of the female seal measured by Gavard was 300 kilos (661 lb.), and that measured by 
Troitzky was 302 kilos (666 lb.) without the viscera. 

There is a certain amount of variation in the colour of the adult seal. Gavard 
says that it is all black except for some patches, particularly a large one of a dirty 


white colour round the navel, and some little yellowish patches on top of the head. 
Agacino (1950) notes that the seal is spotted ventrally, but that there is some 
variation in colour and the old male may be a silvery colour all over. The light 
yellowish beige colour of the female described by Troitzky was also probably due to 
age. Cousteau and Dumas (1953) saw a large white bull in one of the caves on the 
islands of La Galite, but this may have been the silvery colour of old age. The monk 
seal now in the Jardin des Plantes in Paris is thought to be an adult animal and is 
chocolate brown dorsally shading to greyish fawn ventrally. It did not appear to 
have a white ventral patch but had several whitish scar marks along the back. An 
examination was made of the few skins of this species in the British Museum 
collections. The dressed skin of a fairly young male animal, length from nose to tip 
of tail 1 -75 m. (5*74 ft.) (1894.7.27.3, 1063b), is dark blackish brown dorsally, 
with a slightly yellowish appearance due to the yellow tips of the dark brown hairs. 
This shades to light brownish yellow ventrally, but without a light ventral patch. 
The skin of an adult male (length 2-38 m., 1890. 12. 30.1) is generally dark blackish 
brown, slightly yellowish along the centre of the back and belly due to the yellow 
hair tips. In the centre of the belly, slightly nearer the fore flippers than the tail, 
is a roughly diamond-shaped patch of dirty yellow colour, about 73 cm. long and 58 
cm. wide. The whole of the skin, particularly under the chin, the sides of the neck 
and the centre and hinder region of the back, is covered with irregularly placed 
streaks and spots of yellowish hair. These marks show on the under surface of the 
prepared skin and may possibly be due to scars. The skin of the adult female 
(length 2-i m, 1894.7.27.2, io63g) is more like the young animal than the adult 
male. It is dark blackish yellow dorsally, the hairs being dark brown with yellowish 
tips. This shades to light greyish yellow ventrally, and there is no light ventral 
patch. The back shows a few light scar-like streaks similar to those found on the 
male. These whitish streaks were also noticed on the seals examined by Hermann 
and Carruccio (1893) ; they are found on both sexes. The white ventral patch 
seems to be irregular in its occurrence. 

The hairs of the adult are very short and bristly and lie close to the body. They 
are approximately half a centimetre long. The appearance of the young seal 
before it moults its natal coat is described from the youngest skin in the Museum 
collections (1892. 11. 7.1, 1063I). The length of the skin from nose to tip of tail is 
1 -4 m. (4.4 ft.). Dorsally it is a rich dark brown, shading at the level of the fore 
flippers to a lightish brown ventrally. On the belly, slightly nearer the fore flippers 
than the tail, is a roughly diamond-shaped patch of a dirty yellowish colour with a 
few very small light brown spots on it. The patch is approximately 34 cm. long 
and 28 cm. wide at its widest point. There is a small light brown area along the 
upper lip, and the whiskers are also light brown and oval in cross section. The 
texture of the hair of this young seal is quite different from that of the adult. It is 
soft and woolly, the hairs are 1-1J cm. long and do not lie close to the body as they 
do in the adult. 

The whiskers range in colour from light yellow to brown ; they are smooth, not 
wavy as in P. vitulina, and oval in cross section. Nails are present on both fore and 


hind flippers. That on the first digit of the fore flipper is about 2-54 cm. (1 in.) 
long, and the others decrease slightly in size towards the fifth digit. The nails on 
the hind flippers are very small and inconspicuous. The tongue has a notch in its 
anterior end. 

Lobstein (18 17) described in some detail the anatomy of the viscera of a female 
seal that died in Strasbourg in 1815 after touring France and Germany for two years 
and Troitzky (1953) gives a brief description of the viscera of a full term foetus, but 
the most detailed recent investigation into the internal anatomy of M. monachus 
was done by Dieuzeide (1927) on a young male seal that was captured near Oran in 
December 1926, and which lived for a few months in the Experimental Station at 
Castiglione, Italy. A brief summary of the results, taken from Dieuzeide unless 
otherwise noted, is given here for completeness. The length of the male animal was 
1 -68 m. (5-5 ft.) from nose to tip of tail, and of Lobstein's female 2-13 m. (7-0 ft.). 
The oesophagus which lay mainly to the left of the trachea and large blood vessels 
was 60 cm. long and 4 cm. in diameter and was very dilatable. The oesophagus of 
the adult female (Lobstein) was 97 cm. long, the small intestine 14-5 m. and the 
large intestine 1-3 m. The whole alimentary canal was 17-5 m. (5-74 ft.) — eight 
times the length of the animal. There was no definite line of demarcation between 
the oesophagus and the stomach, which looked more like a dilatation of the 
oesophagus. The height of the stomach was about 25 cm., its width about 12 cm. 
and it led into the small intestine, which had a length of 12 m. and a diameter of 
3-3 J cm. The caecum was a small pocket and the large intestine measured a metre 
in length by 4 cm. in diameter. The liver was large, measuring 40 x 40 cm. and 
consisted of six long pointed lobes. There was a large venous sinus (a dilatation of 
the inferior vena cava) and the round ligament (the remains of the umbilical vein) 
was well represented. The gall bladder was multilobulate and when full measured 
13 x 13 mm. The left lung was the larger, weighing 950 gm., while the right weighed 
850 gm. The kidneys were two oval masses, the right anterior to the left, with a 
combined weight of 1 kg. 140 gm. They were lobed and had an extensive vascular 
network. The left renal vein was large and of practically the same diameter as the 
vena cava. It was formed from the union of three large vessels and the superficial 
plexus of the kidney. On the right side the vena cava followed the inner border of 
the kidney and received vessels from it. The right kidney was supplied from the 
aorta by two renal arteries which arose a little anterior to the one going to the left 
kidney. The suprarenals were small and almond shaped, measuring 33 X 14 mm. 
The bladder was very thick (9 mm.) and measured 15 x 4 cm. The prostate was 
reduced and the testes were internal, measuring 4x1.5 cm. The penis was 8 cm. 
long and the baculum 7.8 cm. long. The heart was oval in shape, and Dieuzeide 
gives a detailed description of its anatomy. 

Several species of parasites have been recorded from the stomach and intestine 
(Joyeux and Baer 1936, Baylis 1937, Markowski 1952). There are two Nematodes — 
Contracaecum osculatum, (Rudolphi 1802) and Porrocaecum decipiens (Krabbe 1878), 
and four species of Cestode — Diphyllobothrium coniceps Linstow 1907, D. elegans 
(Krabbe 1865), D. lanceolatum (Krabbe 1865) and D - hians (Diesing 1850). 


Remarkably little information is available about the breeding habits of this seal. 
Practically all that is known is in a recent paper by Troitzky (1953). In this she 
mentions that after a gestation period of eleven months the pups are born on land in 
September and October and are fed by the mother for six or seven weeks. The 
female has four teats on the posterior part of the abdomen and lies on her side when 
feeding the pup. At the end of this time the pup moults its woolly coat and enters 
the water for the first time. Troitzky says that the young seals stay with their 
mothers for three years, that they do not begin to breed until they are four years 
old, and that mating of the adult animals takes place about seven or eight weeks 
after the birth of the pup. The sum of eleven months' gestation plus seven or eight 
weeks before mating makes a total breeding cycle of thirteen months, and although 
it is usual in most Phocids for the female to bear a pup at the same time every year, 
Troitzky says that this seal only has a pup every alternate year. 

Apart from Troitzky's paper, most of the information about breeding times has to 
be inferred from records of still born pups, foetuses and young animals. Dathe 
(1934) records that a young female seal was caught on 19th September, 1933, on the 
Dalmatian coast. The umbilicus was not yet healed and the animal was thought 
to be only a few days old. Its length was approximately 90 cm. and it weighed 26 
kilos. This little animal was kept in captivity and was going to be taken to 
Frankfurt Zoological Gardens. It was fed by bottle six or seven times a day on a 
mixture of half gruel and half milk with a little cod liver oil and freshly rubbed fish 
paste. On 26th September it was taken to Split, preparatory to moving to 
Frankfurt, but the journey re-opened a wound on its stomach, caused probably by 
a fish hook, and on 29th September it died, its length then being 1 . 20 m. 

Carruccio (1893) notes that a foetus 50 cm. long was taken from a pregnant female 
on 21st May, 1891, and Gavard (1927) mentions a captive female that produced a 
still-born pup on 14th April, 1926. This pup weighed 2.25 kilos and measured 62 
cm. Both these foetuses are of such a size that they would have been full term and 
born about September, and it is strange that Postel (1950) should say that the pups 
are born in the spring. Agacino (1950), who went to Las Cuevecillas, Rio do Oro on 
26th December, 1945, says that at that time the smallest seals were 1.5 m. long, 
and that a mother was seen to be feeding her pup. This must have been a pup 
born very late in the season, probably about the middle of November. 

The voice is said to be a sharp strong cry from the bottom of the throat (Cuvier, 
1813), while Agacino (1950) says that when they are annoyed they make a noise like 
a wounded dog. Hermann said that the one he observed had a voice like that of a 
hoarse dog and that sometimes it would howl. This seal could not tolerate dogs, 
and would try to drive them away by clapping its teeth. The seals kept in captivity 
have all been noticeably intelligent and docile animals. They have become attached 
to their keeper and would recognize him, follow him about, and even obey his orders 
to a certain extent. 

The feeding habits, as observed in captivity, are very interesting. The animal 
described by Hermann ate about 14 pounds of fish daily and in order to stress the 
expense of keeping it, inquisitive spectators were told that it ate only the best fish, 


such as eels and trout. It did occasionally receive eels and carp, if paid for by the 
spectators, but usually it had whiting. It took the fish either out of the keeper's 
hands or caught them in its tank, but preferred to eat them in water. It seized the 
fish by the head, squeezed and shook them a few times and then swallowed them 
whole. Often intestines of the fish were found in the water, and although the 
keeper thought this was done deliberately, Hermann suggested that the insides of 
the fish came out accidentally when it was squeezed. Buff on saw the same animal 
while in Paris and said that there it was fed mainly on carp and eels, preferring the 
latter. The fish were sprinked with salt, the eels eaten whole, but the carp were 
crushed with the teeth, let fall and then the belly of the fish ripped open and the 
entrails removed. The fish was then seized by the head and swallowed. Cuvier 
also notes that the entrails were removed and the fish swallowed head first. The 
seal at present in Paris was also seen to swallow its fish head first, and this has also 
been observed in Halichoerus grypus and Phoca vitulina. Indeed it seems possible 
that this method might be the normal one for Pinnipeds generally, as it would avoid 
any injury to the seal by the backwardly projecting fins, scales and spines of the fish. 
Two female seals described by Gavard (1927) also disembowelled their fish unless 
they were very small, and also were unable to pick the fish off the ground, and could 
eat them only in water. They ate sardines, bonito and octopus — about 12 kg. a 
day. Boettger (1951) notes that native fishermen along the African coast say that 
the seals eat fish and lobsters (Palinurus) , and remains of fish of the genera Dentex 
and Labrax have been found in the stomach of a seal captured off Sardinia (Carruccio 
1893). A monk seal in the Gulf of Salonika was seen to be playing with a large fish, 
tossing it into the air and catching it again (Butler in litt.) 

Troitzky (1953) describes an adult female seal that was killed in a cave on Corsica 
in September, 1947. It was found to be pregnant and a full-term foetus was 
removed, but could not be revived. The pup, a female, was 120 cm. long and 
weighed 17 kilos. It was dark, greyish black colour, with a white ventral patch. 
Troitzky notes that the pup in its colouring did not differ from descriptions of what 
she regarded as " typical " M. monachus, but says that the mother was not so. 
She observes that in its dentition, its great size, and the time of breeding the adult 
resembles M. monachus, but the shape of its head with a long snout, the light colour, 
the absence of the white ventral patch, and the second digit of the fore flipper longer 
than the first are characters not associated with that species. After reviewing other 
members of the Phocidae she comes to the conclusion that these characters are more 
like those of Arctic Phocids, and says that the most logical conclusion is that this 
female is a hybrid, the result of a cross between a monk seal and, probably, Phoca 
groenlandica, and that it is interesting that such a hybrid should have been able to 
produce a pup. 

It is considered extremely unlikely that such a cross could have taken place. 
Phoca groenlandica and other members of the Arctic Phocidae have not been recorded 
from the Mediterranean in Recent times, and indeed, they seldom occur south of 
Arctic latitudes. From the description of the teeth — worn, broken and diseased — 
it is assumed that the animal was old, and the light colouring was probably also due 



to age. The white ventral patch is not of universal occurrence amongst the monk 
seals. It is not possible to comment on the other two characters, from lack of exact 
information, but from the photograph of the adult seal, as far as it is possible to see 
the shape of the fore flipper does not appear to be unusual and the drawings and 
photograph of the skull do not appear to differ in any way from undoubted skulls of 
M . monachus in the Museum collection. 

II. Monachus tropicalis 


The first reference to the monk seal of the West Indies is that given in the account 
of the second voyage of Columbus. At the end of August, 1494, the ship anchored 
by the rocky island of Alta Vela, south of Haiti (= Hispaniola), and the men that 
went ashore killed eight " sea wolves " that lay sleeping on the sand (Kerr, 1824). 
The next record chronologically, and the first for Florida, is that of Herrera who, 
while describing Ponce de Leon's discovery of the Dry Tortugas Islands (lat. 24 10' 
N. long. 83 55' W.) on 21st June, 1513, said that a foraging shore party took 
fourteen seals (Moore 1953). Du Tertre (1667) was told by Brother Charles Poncet, 
who had been to Guadeloupe, that he had seen at least twenty asleep under the trees 
near the shore, and many of them were killed. Dampier (1705) noted that there 
were seals on the Alacrane Islands in 1675, Sloane (1707) saw them on the Bahama 
Islands in 1687, and Olafsen (1774) makes a reference to the seal of the Antilles. 
Hill (1843) gives a description of a young seal from Pedro Kays, and Gosse (1851) 
published an account of a voyage in 1846 by Mr. George Wilkie to Pedro Kays, 
where he saw several seals and killed a few. It was on this voyage that the type 
skin of Phoca tropicalis was obtained. In 1883 the U.S. National Museum received 
a mounted skull and skin from a female seal captured off Cuba earlier in the same 
year and presented by Prof. Felipe Poey (True and Lucas, 1885). In 1886 H. L. 
Ward and Prof. F. Ferrari Perez of the Mexican Geographical and Exploring Survey 
set out from Campeche to the Triangles to search for Monachus. (Ward, 1887b). 
Although they were only on the islands from 1st to 4th December, forty two 
specimens were taken away and shared between the two members of the expedition. 
Of the specimens retained by H. L. Ward a complete articulated specimen (1887 . 8 . 
5.1), and a skin and skull (1889. n. 5.1) are now in the British Museum (Natural 
History), and an articulated specimen (899c) is in the Cambridge Zoological Museum. 
These last two specimens were purchased from Ward by F. D. Godman of Cambridge. 
A recently born pup was taken back to Campeche, but it lived there only a week. 
A female seal was captured at the Triangles in 1897 and lived for nearly 5 J years in 
New York Aquarium, where it died in 1903. (Anon 1903). E. W. Nelson and E. A. 
Goldman, during their biological investigations of Mexico from 1892 to 1906 
(Goldman, 1951), spent the period 18th to 23rd June, 1900, on the Triangles. Their 
main object there was to obtain specimens of seals, but apart from remarking that 
" in quest of these animals we were very successful ", no mention is made of their 
abundance, or how many were killed. The New York Aquarium received four more 


seals — an adult male and three yearlings — in June, 1909 (Townsend 1909). These 
were obtained from a dealer in Yucatan who presumably got them from from the 
Triangles or the Alacrane Islands. Several seals were seen on the Tortugas Islands 
during the period 1903-8 and two were kept in a moat for some time, where they 
became fairly tame (Moore, 1953). Six seals were captured by a fishing vessel in 
1915 and taken to Pensacola, where they were kept in captivity for some time and 
then turned loose, when bathers in the area objected to their presence (Allen 1942). 
Townsend (1923) notes that a seal was killed near Key West, Florida in March, 1922. 
Gunter (1947) gives sight records of seals along the Texas coast in 1926 and 1932, 
and Lewis (1948) says that a young seal was killed at South West Kay in the Pedro 
Group in 1939. A. C. Wheeler (in litt.) saw two seals on the beach of Drunken 
Man's Cay, about two miles south of Kingston, Jamaica in November, 1949. 

Distribution and abundance 

The West Indian monk seal was at one time abundant in the Gulf of Mexico and 
off the islands in the Caribbean Sea (Fig. 3.). The presence of numerous Seal Cays 
and Islands and Lobos Cays show how widely the seal was distributed in the area. 

1. Seal Cay, south of Long Island, Bahamas, lat. 22 38' N., long. 75 ° 54' W. 

2. Seal Cay, south of Caicos Bank, Bahamas lat. 21 10' N, long. 71 47' W. 

3. Seal Island, north of Anguilla, Leeward Is, W. Indies lat. 18 24' N., long. 

4. Cay Lobos, north of Cuba, lat. 22 25' N., long. yy° 36' W. 
and Allen (1887a) gives the following : 

5. Seal Keys — on the coast of Honduras in about lat. 16 N., a few miles north- 
east of the Mosquito Coast. 

6. Seal Key — about 200 miles further south along the same coast in about 
lat. 12 40' N. 

It has been recorded from the Bahama Islands ; Key West, Florida ; Galveston 
Bay and Brazos, Texas ; the Triangle Islands to the west of Yucatan, lat. 20 55' N. 
long. 92 12' W. : the Alacrane reef to the north of Yucatan, lat. 22 32' N., long. 
89 45' W. ; the shores of Cuba and Jamaica ; the Pedro Kays to the south of 
Jamaica, lat. 17 N., long. 77 30' W. ; Alta Vela, a rocky island south of Haiti ; 
and Guadeloupe. 

That the seal has formerly been abundant is evident from some of the earlier 
accounts. Dampier (1705) said of his visit to the Alacranes in 1675 : " Here are 
many seals . . . the Spaniards do often come hither to make Oyl of their Fat ; 
upon which account it has been visited by English-men from Jamaica, particularly 
by Capt. Long : who having the Command of a small Bark, came hither purposely 
to make Seal-Oyl." Sloane (1707) gives the following account of the seals of the 
Bahamas, " The Bahama Islands are fuTd with Seals, sometimes Fishers will catch 
one hundred in a night. They try or melt them, and bring off their Oil for Lamps 
to these Islands." In 1856 a Mr. Alexander was on the Triangles and saw only two 










ZOOL. 3, 5. 



living seals, but remains of skeletons and hides indicated a once flourishing business ; 
and although H. L. Ward does not say exactly how many he saw in 1886, there must 
have been quite a large colony as he killed over forty animals (Ward, 18876). Allen 
(1887a) suggested that at the time of his writing seals were still present on the islets 
of Salt Key Bank, north of Cuba, the isles off Yucatan, and probably the isles 
between. In 1897 at the time of the capture of the seal for New York Aquarium 
about thirty were observed on the Triangles (Anon 1903), but as late as January, 1911, 
according to Lewis (1948) about two hundred seals were killed in this locality and at 
the time his paper was published he thought that there were perhaps still a few left 
there and on the South West Kay in the Pedro Group. In view of the fact that 
seals have been seen as recently as 1949 near Jamaica, and that Moore (1953) quotes 
a " well informed and responsible friend " who knows of the whereabouts of seals 
somewhere within their former range, it seems likely that a remnant of this species 
is still living. 


During Mr. George Wilkie's visit to the Pedro Kays in 1846 (Gosse, 1851) he 
obtained the skin of a monk seal which he gave to Gosse. Gosse sent this skin, which 
had no bones with it, to the British Museum where J. E. Gray published a description 
of it under the name Phoca tropicalis Gray, 1850. This skin has been stuffed and is 
in the Museum collection (Reg. No. 1847.2.2.2). Gosse (1851) republished the 
description of the seal given by Hill (1843), and in a footnote on p. 308 suggested 
the specific epithet " wilkianus ", but he gave no generic name, and moreover, was 
already antedated by Gray. In 1866 in the Catalogue of the Seals and Whales in the 
British Museum Gray repeated the descriptive paragraph he gave in 1850, but used 
the name Monachus tropicalis. 


The nose to tail length of an adult male animal is between 7 and 8 ft., females being 
in general slightly smaller. Townsend (1906) gives the length of an old female with 
very worn teeth as 9 ft., but he was probably measuring to the tips of the hind 
flippers. The nose to tail lengths of both the skeleton in the British Museum, 
(probably a female) and that in Cambridge, (sex unknown) are both about 7 ft. 3 in. 
The nose to tail length of a dressed skin of an adult male (1889. 11. 5.1, 1064b) is 
7 ft. 5 in., and Ward (18876) gives nose to tail lengths of two pregnant females as 7 ft. 
1 in. and 6 ft. 6 in. A female measuring 6 ft. 11 in. to the tip of the hind flippers 
and which had been in New York Aquarium for 5^ years weighed 360 pounds at 
death. This seal died from fatty degeneration of the heart, liver and kidneys, so 
presumably a healthy animal of this size would weigh less. 

Ward (18876) gives the following description of the colour of the adult seal : 
" Adults are grayish brown or grisled on the back, a result of the Vandyke-brown 
hairs being tipped with light horn-color, the lower surface ochreous yellow to 
yellowish white. Females seem to have much less of the yellow or white on the 
ventral surface. From the black pelage of the extremely young to that of the adult 



there is an intermediate stage of yellowish gray on the dorsal surface, shading to 
almost a perfect ochre on the ventral portions." This is the most reliable account of 
the colour, and corresponds well with that of the stuffed specimen in the Museum, 
which is the type of Phoca tropicalis Gray. This animal is dark brown, and slightly 
lighter on the sides and belly. The hairs of the back are dark brown with a lighter 
tip, while those of the sides and belly have a more yellow tip. The hairs lie close to 
the body and are extremely short, the longest — those on the sides — being about 1 cm. 
long. A dressed skin of an adult male from the Triangles in the Museum collection 
(1889. 11. 5. 1, 1064b) is dark blackish brown all over, with a slight yellowish tinge 
due to the yellow tips of the hairs. New born pups are black in colour and the hair 
is long, soft and woolly. 

The whiskers are yellowish horn colour, some being slightly darker at their bases. 
Those of the Museum specimens are oval in cross section. Nails of appreciable size 
are present only on the fore flippers. That on the first digit is about an inch long, 
those on the remaining four digits decreasing gradually in size. The nails of the 
hind flipper are very small. 

The description of the eye is given by Ward (18876) : " The pupil is medium 
sized, round, and well defined, the iris is light reddish brown in color, and with but 
little of the sclerotic coat showing. Over the cornea there appears a deadening 
film . . .", which he attributes to the strong reflection of the tropical sun from the 
coral sands. 

Young animals are born about the beginning of December. Ward (18876) was on 
the Triangles from 1st to 4th December and killed five females with full-term 
foetuses, and noticed another female with a new-born pup. The female has four 
teats. One of the foetuses measured 85 cm. from nose to tip of tail, was covered in 
black woolly hair and had black whiskers. Measurements of the skull and skeleton 
of this pup as well as those of adult animals collected by Ward are given by Allen 

Ward examined the stomach contents of several animals, but found only fluids 
and large numbers of intestinal parasites several inches in length. Gosse (1851) 
recorded the opinion of the " more experienced fishermen ", who said that the seals 
fed " as generally on molluscous animals as on fish ", but there is no proof of this 
The animal noted by Hill (1843) lived for four months in captivity, without eating 
and when it died " the fat was four inches thick and yielded four gallons of oil " 
The skull of this animal, which was then, as Hill (1846) says " an undescribed Seal " 
was exhibited at a meeting of the Zoological Society of London in September, 1846 
Unfortunately it has not been possible to trace this skull, which was probably the 
first specimen of the West Indian monk seal to reach this country, arriving here 
shortly before the skin sent by Gosse which became the type. 

The West Indian monk seal appears to be a fairly noisy animal. Hill (1843) said 
that his young animal " grunted, barked, growled and snarled like a dog ", and 
Ward (18876) said that the voice of the young was " a long drawn out guttural ' ah ' 
with a series of vocal hitches during its enunciation ". Townsend (1909) also noted 
that it was noisy, and the young often roared harshly. 


On land at least the seal seems to be rather lethargic. Ward (18876) notes that 
unless the seals were approached closer than three or four feet they showed no 
interest or alarm. Closer than that they would rouse themselves, bark, and move 
off a little. When Ward and his party attacked a group of seals they got more 
excited and would make savage rushes, and would then fall back on their dead 
fellows and bite and shake them. Nevertheless, as Ward notes " the whole aspect 
of the animals was one of indecision . . . they only roused themselves to action 
on being individually attacked ". This behaviour is not peculiar to the monk seal, 
but seems to be common to all seals. Ward also notes the peculiar circumstance 
that several of the animals he collected had a growth of minute algae upon their 
backs and flippers, especially the hinder ones, so that they appeared quite green. 

Although Ward says they are neither curious nor playful, Hill (1843) notes that 
the young specimen he kept in captivity was lively, and those kept in New York 
Aquarium were certainly playful. The two which were received in 1897 had the 
habit of filling their cheeks with water and squirting it at visitors, while the seal 
which arrived in 1909 amused itself by tossing flipperfuls of water into the faces 
of visitors. 

III. Monachus schauinslandi 


The Hawaiian or Sandwich Islands are a chain of small islands near the centre of 
the North Pacific Ocean between 18 55' and 28 25' N., and 154 48' and 178 25' W. 
(Text-fig. 4) ; Honolulu on the island of Oahu being 2,100 miles S.W. from San 
Francisco and 3,445 miles S.E. from Yokohama. The chain stretches 1,578 miles 
from E.S.E. to W.N.W. All the islands are uninhabited, except Midway which is a 
transpacific cable station and sea plane base administered by the U.S. Navy, and 
Niihau, Kauai, Oahu, Molokai, Maui and Hawaii. The entire chain forms the 
Hawaiian Islands Bird Reservation. 

Perhaps owing to the remoteness of the islands on which they live, there are very 
few references to the Laysan monk seal. In the early part of the nineteenth century 
seals must have been numerous as Bryan (1915) records that in 1824 the brig 
" Ainoa " set out from Hawaii on a sealing voyage in that area, and in 1859 * ne 
" Gambia " returned to Honolulu with 1,500 seal skins and 240 barrels of seal oil, 
some of which was probably from Midway Id., which was discovered on this trip. 
In 1893 a Mr. J. J. Williams visited Laysan and heard of an earlier expedition that 
had killed sixty or seventy seals on the island (Atkinson and Bryan, 1913). In 1905 
Matschie published a description of a seal skull brought back from Laysan by Dr. 
H. Schauinsland, and named it after him. The U.S. revenue cutter " Thetis " 
returned in 191 2 after a cruise to Midway and Laysan and brought back a seal skin 
which was presented to the Bishop Museum in Honolulu (Bryan, 1915) and parts of 
three others which are in The U.S. National Museum. Thirty five seals were seen 
on Pearl and Hermes Reef in 1913 (Atkinson and Bryan 1913), and Dr. Wetmore, 
who visited the area with the U.S.S. " Tanager " Expedition in 1923 saw a number of 
seals and collected ten for the U.S. National Museum (Bailey, 1952). In 1940 about 













CD Tt 














or ^ 










"V 3" 
S ft* 






half a dozen were seen round Midway (Blackman, 1941) and records of visits to the 
islands in 1949 and 1951 show that quite large numbers were seen resting on the 
beaches. The first, and it is believed, the only seal of this species to be kept in captivity 
outside the Hawaiian Islands was a young animal which was presented to San Diego 
Zoo by the Honolulu Zoo in 1951 (Anon, 1951). It was captured on French Frigates 
Shoal but only lived about three months in captivity. 

Distribution and abundance 

The Laysan seal has been recorded from the following of the Hawaiian islands : 
Ocean Island, Midway Island, Pearl and Hermes Reef, Lisiansky Island, Laysan 
Island, French Frigates Shoal, and a stray animal has been recorded from the coast 
of Hawaii, although they do not generally appear to go so far eastwards. 

In 1824, x ^59 an d 1893 the seals were obviously very numerous, although in 
Schauinsland's own account of his visit to Laysan (Schauinsland, 1899) ne OIU y 
mentions that " seals come singly, indeed very seldom by the island ". In 1912, 
Elschner (1915) who was also on the "Thetis" during its trip, noted that there were 
many seals on French Frigates Shoal and Pearl and Hermes Reef. Also on the latter 
island thirty five seals were seen in 1913 (Atkinson and Bryan, 1913). The U.S.S. 
" Tanager " Expedition sailed from Honolulu in April, 1923, for a visit to the Hawaiian 
Islands. The Bulletin of the Bishop Museum in Honolulu (Bull. 10, 1924) only 
mentions that several seals were seen on Lisiansky on this voyage, and that two 
skins and skulls were collected ; Allen (1942) however, notes that Dr. A. Wetmore 
while on this expedition saw colonies of seals on Pearl and Hermes Reef and on Ocean 
Island, and estimated the total population to be about four hundred, and Bailey 
(1952) notes that he brought back ten specimens for The U.S. National Museum. 
Sixty eight seals were seen on Pearl and Hermes Reef in 1930, and five on Laysan in 
1936 (Bailey, 1952), and Blackman (1941) who stayed for six months on Midway in 
1939-40 saw about six seals. More recent visits to the islands (Bailey, 1952) show 
that the animals are still fairly numerous there. About thirty were seen on Laysan 
and over a hundred on Midway. A count of seals made in 1951 showed a total of 
407 on the beaches, the largest populations being on Laysan (119) and Pearl and 
Hermes Reef (180). The others were seen on French Frigates Shoal, Maro Reef, 
Lisiansky and Midway. It was suggested that the large numbers on the beaches 
could perhaps be accounted for by the presence of tiger sharks in the water. 

The monk seals are distinctive in being the only truly warm water form of Phocid, 
with the exception of the northern elephant seal (Mirounga angustirostris) . So far 
as the distribution of the three species is concerned it is not difficult to accept the 
occupation of the West Indian islands from a source in the Mediterranean and along 
the Mauritanian coast, as the Canary Current passing down the latter coast would 
bear the emigrants to the eastern limit of the North Equatorial Current sweeping due 
west to the Caribbean Sea. The extension of the range of the genus to Hawaii and 
across the Isthmus of Panama is feasible when it is accepted that Phocids are capable 
of considerable overland journeys. For example Hayes (1928, p. 106) records that 
Captain Scott, when in the Antarctic, found seal carcases as far as fifty miles inland 


and 5,000 ft. above the sea. The Isthmus of Panama at its narrowest is much less 
than fifty miles, and its lowest height above sea level less than 200 ft. If rivers 
were exploited by the seals the distances travelled overland might have been still 
further diminished. The North Equatorial Current in the Pacific could well have 
borne the animals to the islands they now occupy. Allen (1942) suggests that the 
colonization of the Pacific by the monk seal was from the West Indian stock in 
Tertiary time, when there was a waterway connecting the Atlantic and Pacific. 


Of the few records available only those of Matschie and Bailey (1952) give any 
information about the external features and Bailey includes some excellent photo- 
graphs. A male, probably not full grown, collected by the " Thetis " measured 5-7 
ft. from tip of nose to tip of tail, and a female 7-5 ft. Matschie gives the following 
description : "A scalp which I have for examination has very short, thick, bristly 
hair, loam coloured, with a silky shine. The lips are yellow-gray, and it is somewhat 
brownish on the front side of the neck. Most of the whiskers are light horn coloured, 
a few dark brown, and all with very thin points. The sides of the body of the stuffed 
animal in Bremen Museum are lighter than the back, the breast and stomach are 
whitish." Bailey notes that the underparts are light straw yellow and the back is 
dark slate grey. 

While on the " Tanager " Expedition Dr. Wetmore examined the stomachs of 
seals for parasites. All the seals had abundant nematodes in the stomach and 
Chapin (1925) has described these as a new species Contracaecum turgidum. 
Cestodes, Diphyllobothrium Mans, were also found. 

The birth of the young seals evidently takes in January. The " Thetis " saw 
pups at this time in 1912, and one was born during the visit of Governor Frear to 
Pearl and Hermes Reef in January, 1913 (Atkinson and Bryan, 1913). The young 
male seal that lived in San Diego Zoo for a short time (Anon, 1951) was received in 
May and died probably at the beginning of September. It was therefore about eight 
months old at the time of its death. It is described as a nursing pup of approximately 
three feet long. It had a silver-tinged coat which was dark brown above, paler on the 
sides and nearly white ventrally. The muzzle was whitish and had many coarse 

Atkinson and Bryan note that the seals are fearless and readily handled, and 
Blackman (1941) says that if cornered they threaten the intruder by opening their 
mouths widely and uttering an abrupt barking noise. He also says that they do 
not migrate, and probably feed on squid and fish. Bailey notes that an animal 
collected by Henry Palmer in 1891 had its stomach full of half digested fish ; and he 
also makes an interesting observation that a large male seen in 1949 had a greenish- 
coloured face, but what the cause of this was he did not say. 


The following specimens of Monachus have been examined. The list includes 
stuffed animals and skins, and unless otherwise mentioned the specimens are in the 
British Museum (Natural History) . 


M. monachus 

Skulls only 

i. 1853.10.6.4,1063a. Presented by R. Macandrew. 
Deserta Grande Is. Madeira. 
Type of Heliophoca atlantica Gray, 1854. 
Skull and lower jaw of immature animal. Skull with the dorsal part of the 

cranium, inter orbital and nasal regions missing. 
Stuffed animal also in the collection. 

2. 1063b. Presented by R. Macandrew. 
Deserta Grande Is. Madeira. 

Skull incomplete — maxillary region bearing teeth, and fragmentary lower jaw 

3. 1934.8.5.4 Collected by Barrett-Hamilton. 

Complete skull and lower jaw. 

4. 1951 . 4 . 17 . 1. From Rothschild Collection, Tring. 
Skull and lower jaw. Zygomatic arches broken. 

Skulls and skeletons 

1. 1063c. Presented by M. Verreaux. 

Complete skull and skeleton of an immature animal. 

2. 1863.4.1.1,1421a. Male. Purchased from an Italian. 
N. Mediterranean. 

Complete skull and skeleton of an immature animal. 

3. 1892. 10.4. 1, 1063d. Presented by C. F. R. Blandy. 
Deserta Grande Is. Madeira. 

Complete skull and skeleton of an immature animal. 

4. 1892. 11. 7. 1, 1063I. Presented by C. F. R. Blandy. 
Deserta Grande Is. Madeira. 

Complete skull and skeleton of very young animal. 
Skin also in collection. 

5. 1894.7.27. 1, 1063! Male. Presented by C. F. R. Blandy. 
Deserta Grande Is. Madeira. 

Complete skull and skeleton of adult animal. 

6. 1894.7.27.2, io63g. Female. Presented by C. F. R. Blandy. 
Deserta Grande Is. Madeira. 

Complete skull and skeleton of adult animal. 
Skin also in collection. 

7. 1894.7.27.3, 1063I1. Male. Presented by C. F. R. Blandy. 
Deserta Grande Is. Madeira. 

Complete skull and skeleton of immature animal. 


The seal was sent alive from Madeira and lived in the Zoological Gardens 

London from 16th July to 28th October, 1894. 
Skin also in collection. 
8. 1951.4.17.2. No history. 

Incomplete skeleton of an immature animal. 
No skull. 


1. 1890. 12. 30. 1. Male. Presented by H. C. Hinton and C. J. Cossart. 
Bugio, Deserta Grande Is. Madeira. 

Skin of an adult animal. 

2. 1892. 11. 7. 1, 1063I. 

Skin of very young animal noted above. 

3. 1894.7.27.2, io63g. Female. 
Skin of adult animal noted above. 

4. 1894.7.27.3, 1063I1. Male. 

Skin of immature animal noted above. 

Stuffed animals 

1. 1853. 10.6.4, 1063a. 

Belonging to type of H. atlantica Gray, noted above. 

2. 1 910 . 9 . 27 . 1. Presented by Zoological Society of London. 

Presented to Zoological Society by Godfrey Williams. 

3. An immature animal with no history. 

M. tropicalis 

Skull only 

1. 1889 .11.5.1, 1064b. Male. Presented by F. D. Godman. 
Triangle Is. Gulf of Mexico. 

Collected in 1886 by Comision Geografico Exploradoro, Mexico. 
Skull and lower jaw of adult animal. Skull complete except for both jugals. 
Skin also in collection. 

Skull and skeleton 

1. 1887 .8.5.1. Collected by H. L. Ward. 
Triangle Is. Gulf of Mexico. 

Complete skull and skeleton of adult, probably female animal. 

2. K.7801, 899c. Presented by F. D. Godman. 
Triangle Is. Gulf of Mexico. 

Complete skull and skeleton of adult animal. In University Zoological 
Museum, Cambridge. 



I. 1889. 1 1. 5. i, 1064b. Male. 

Skin of adult animal noted above. 

Stuffed animal 

1. 1847.2.2.2. Presented by P. H. Gosse. 
Type of Phoca tropicalis Gray, 1850. 

M. schauinslandi 

i- 32795. 

The skull of the type specimen collected on Laysan Id. by Dr. Schauinsland. 
The skull is in the Zoological Museum in Berlin and has not been examined, 
but photographs have been made available by Dr. K. Zimmermann, and a 
very complete set of measurements was published by Matschie (1905). 

A stuffed specimen is believed to be in the Bremen Museum. 


A skull belonging to the genus Monachus has the following characters : 

1. The skull is broad in proportion to its length. 

2. The dorsal surface is convex, sloping backwards and forwards from a point 
about halfway along the interorbital region. 

3. The interorbital region is broad and parallel-sided. 

4. The naso-maxillary region is flattened dorsally. 

5. The snout is broad, the sides of the maxillae being almost parallel from the 
upper edge of the infra-orbital foramen to the canines. 

6. The molars are large and set more or less obliquely, and there are four upper 
and four lower incisors. 

7. The condyle of the lower jaw is very low, on a level with a line drawn through 
the points of the molar teeth. 

Comparison of the skulls of M. monachus, M. tropicalis and M. schauinslandi 

1. In view of the small number of skulls examined, few general conclusions can be 
drawn from the measurements and proportions (Table I), although, as the following 
summary shows, skulls of M. monachus tend to be slightly wider in proportion to 
their length than skulls of M. tropicalis and M. schauinslandi (Text figs. 5 and 6, 
PL 6). 

M. monachus. M. tropicalis. M. schauinslandi. 

0/ 0/ 0/ 

/o /o /o 

Zygomatic width . . . . 59- 9-70 -3 . 61-7-62-1 . 61-5 

Snout width at canines . . 20-9-26-0 . 20-6-20-9 . 20-3 

Width at external auditory meatus 53*1-58-5 . 49*8-50-5 . 50 • 5 

Width at petrous bones . . 60 • 3-64 -9 . 56 • 3-59 -2 . 59-2 



10 CMS. 

10 CM5. 

Fig. 5. — M. monachus Reg. No. 1894.7.27. 1. a. Dorsal view of skull, b. Ventral view. 
Note supernumerary incisors, c. Lateral view of skull and lower jaw. 



10 CMS. 

10 CMS. 

Fig. 6. — M. tropicalis Reg. No. 1887.8.5. i. a. Dorsal view of skull, b. Ventral view, 
c. Lateral view of skull and lower jaw. 


2. Each nasal of M. monachus ends anteriorly in a V, the point of the V, which 
may be slightly rounded, being directed anteriorly, so that the ends of both nasal 
bones together form a W. The nasal septum may project slightly beyond the nasal 
bones in old specimens. The nasal bones of M. tropicalis are much longer and 
narrower and their anterior ends continue the curve made by the premaxillae. The 
nasal septum forms a triangular projection. The nasal bones of M. schauinslandi 
are as long as those of M. tropicalis, but do not taper to so fine a point posteriorly. 
The anterior end of each has the form of an inverted V with the point directed 
posteriorly. The nasal septum appears to project a little beyond the end of the 
nasal bones (Text-fig. 7). 

3. When seen from in front, the lower edge of the infraorbital foramen is wider 
than the upper edge in M. monachus, while the reverse is true in M. tropicalis. The 
foramen is not visible in the photographs of M. schauinslandi, although Matschie 
(1905) says that the lower edge is wider than the upper. 

4. There is a well defined tubercle on the maxilla at the anterior margin of the 
orbit in M. monachus. This is very small and indistinct in M. tropicalis, and 
Matschie (1905) says that M. schauinslandi also has no clear tubercle in this position. 

5. In M. monachus the zygomatic branch of the squamosal is not expanded at its 
anterior end and lies at an angle of approximately 45 ° with the base of the skull. 
The orbital process of the jugal is wide and upwardly directed while the lower branch 
is narrower and continues back along the squamosal for some distance. 

In M. tropicalis the zygomatic branch of the squamosal is expanded at its anterior 
end and is much more nearly vertical. The orbital process of the jugal is very 
narrow and curves backwards over the top of the zygomatic branch, while its lower 
branch is short and triangular. 

In M. schauinslandi the zygomatic branch of the squamosal is similar to that of 
M. monachus ; it is not expanded and lies at an approximate angle of 45 °. The 
orbital process of the jugal is not quite so wide as that of M. monachus, but is 
otherwise similar, and the lower branch is long and backwardly directed (Text- 
fig- 7). 

6. The posterior end of the palate of M. monachus is U-shaped with a small median 
V-shaped incision. In M. tropicalis it is V-shaped, and in M. schauinslandi it forms 
a slightly wider V. (Text-fig. 7) . 

7. The pterygoid bones of M. monachus are low and ill-defined, and are not visible 
when the skull is viewed dorsally. In M. tropicalis they curve widely outwards 
and are visible dorsally. In M. schauinslandi they curve out as in M. tropicalis, 
but are not visible dorsally. 

8. The coronoid process of the lower jaw of M. monachus is wide and concave 
internally. That of M. tropicalis is narrow and only very slightly concave. The 
coronoid of M. schauinslandi is narrow and like that of M. tropicalis. 

9. From the above characters it will be seen that the skull of M. schauinslandi is 
more like that of M. tropicalis than M. monachus. 







Fig. 7. — Nasal bones, zygomatic arch and hind end of palate of a. M. monachus, b. M. 
tropicalis, c. M. schauinslandi. sq., squamosal ; ju., jugal. 



Table I. — Cranial Measurements of Monachus Skulls. 

M. monachus. 


4. 17. 1. 






























Condylobasal length . 



. 281 


• 273 


. 268 


. 262 


Condylobasilar length 



. 270 


. 262 


• 258 




Basal length 



• 263 


. 251 


• 251 




Basilar length . 



. 252 


. 241 


. 241 




Snout width at canines 



. 68 


• 7i 


• 57 

21 -2 

■ 55 


Width of skull at front 



. 76 


• 85 


. 60 




end of last upper 


Zygomatic width 




• 193 


. 185 


. 176 

6 5 -6 



Width at upper edge 



. 156 


. 150 


• 145 




auditory meatus 

Width at petrous 


60 -3 

• 177 


. 165 

60 -4 

. 167 


. 160 

61 • 1 


Palatal length . 



• ii7 


• 113 


. no 

41 -o 



Palatilar length 



. 107 


■ 103 






Width of occipital con- 



• 67 


. 64 


. 65 


■ °5 



Length of nasal suture 



. 46 


• 52 






Length of upper molar 


21 -6 

. 60 


. 64 



21 -2 




M. monachus. 


10.4. I. 






II .7.1. 











> k 

> » 













Condylobasal length . 



. 224 


. 217 






Condylo-basilar length 



. 216 


. 212 





95 '9 

Basal length 





. 197 



91 -2 



Basilar length . 





. 191 






Snout width at canines 





. 5i 






Width of skull at front 



• 63 


• 63 






end of last upper 


Zygomatic width 






59*9 ■ 


63-2 . 



Width at upper edge 






56-7 • 


58-5 • 



auditory meatus 

Width at petrous 






62-7 . 


64-9 . 




Palatal length . 






41-9 . 


42-7 . 



Palatilar length 






39*2 . 


38-0 . 



Width of occipital con- 





. 62 

28-6 . 


31-6 . 




Length of nasal suture 






— . 


22-2 . 



Length of upper molar 





• 52 

23*9 • 


26-9 . 




2 32 


Table I — cont. 

M. tropicalis. 












-* ^ 
















Condylobasal length 









Condylo-basilar length . 






96-6 . 



Basal length 









Basilar length 









Snout width at canines 









Width of skull at front end o 

f 77 








last upper molars 

Zygomatic width . 






61 -7 



Width at upper edge auditors 

T I36 









Width at petrous bones 





■ 151 




Palatal length 









Palatilar length . 









Width of occipital condyles 

■ 67 








Length of nasal suture . 

• 67 








Length of upper molar row 



• 65 




M. monachus 



Dental formula : . 2 

1 5 

1-, c 


i' m 

Upper. — The incisors are large and pointed, the point directed posteriorly. The 
outer incisors are larger than the inner and there is a well developed cingulum on the 
inner surface of all four teeth. 

The canines are of moderate size with a slight ridge down the posterior surface. 

The molars are large, the 3rd being the largest and the others decreasing in size in 
the order 3, 2, 4, 1, 5, except in the largest skull (1951.4.17.1) where the 5th is 
slightly greater than the first, and the others decrease in size from before backwards. 
Although the molars are set obliquely they do not overlap one another. The first 
molar makes an angle of 20 with the median palatal suture, the 2nd molar 40 , the 
3rd molar 6o°, the 4th molar 70 and the 5th molar no°, so that its palatal surface 
is facing posteriorly. The external edge of the palate forms an angle of 20 with 
the median palatal suture. These measurements are from 1892.10.4.1, and are 
compared with a Phoca vitulina 1919.7.7.3260, where the teeth are set in line at an 
angle of 20 . There is a slight obliquity in the teeth of some of the other P. vitulina 
skulls, but this is not nearly so pronounced as in M. monachus. All the molars are 
double rooted except the first. The molars have a large central cusp and single 


smaller anterior and posterior cusps. The small cusps on the 5th molar are less 
distinct and the posterior one may be lacking. There is a well developed cingulum 
on the inner surface. 

Variation. — In the nine specimens with fully erupted teeth, variations in the 
number of upper incisors occur in five of them. 

1. 1853. 10. 6. 4, 1063a. 

There is a small extra tooth just posterior to the first left incisor. 

2. 1063b. 

Although the tooth is missing in the specimen, the alveolus shows evidence of an 
extra tooth posterior to the first right incisor. 

3. 1894.7.27. 1, 10631. 

On the right side there is a small incisor median to the first and in the same line 
with it. The first incisor is the same size as that on the left side, though 
farther from the mid line. The second incisor on the right side is separated 
from the first by a gap, and leans away from it, towards the canine. It is 
considerably smaller than the left second incisor. Posteriorly and internally 
to the first and second right incisors is part of the root of a tooth larger than 
the left second incisor and smaller than the canine. The crown has been 
broken off since the animal died. 

4. 1934.8.5.4. 

On the left side a small incisor is set directly behind the first. 

5. 1951.4.17.1. 

On the left side a small incisor is situated just posteriorly to the first. 

Disease. — Only one specimen shows any sign of disease in the teeth. 1863 .4.1.1, 
1421a — the second right incisor has lost most of the crown, the tooth is hollow, and 
there is some lumpy growth on its anterior surface. The parts of the premaxilla 
round the tooth have fallen away. 

Lower. — The first incisors are considerably smaller than the second, they are set 
internally and posteriorly to them and are in a more or less recumbent position. 
They are similar to the upper incisors in shape ; the canines are also similar. 

The third molar is the largest, the others decreasing in size in the order 3, 2, 4, 5, 1, 
so that the 1st molar is the smallest. The molars are set obliquely and do not over- 
lap one another. The first molar is set directly behind the canine, so close that the 
two alveoli are confluent, but the molar is not on the inner surface of the canine as 
in the upper jaw. The angle the teeth make with the symphysial line is not so 
varied as in the upper jaw. The 1st molar makes an angle of 30 , the 2nd 40 , the 
3rd 40 , the 4th 6o°, and the 5th 20 . As the ramus itself is at an angle of 20 the 
5th molar is in line with the jaw. These measurements are from 1863 .4.1.1, 1421a. 
All the molars are double rooted except the first. The shape of the lower molars is 
similar to that of the upper, except that the posterior cusp on the 5th molar is 
generally present. 

There are no variations in number, and no disease is present. The surface of all 
the teeth, both upper and lower, is slightly rugose. 

ZOOL. 3, 5. 17 


M. tropicalis 

Dental formula : . 2 1 5 

l i- c ? m 5 

Upper. — The incisors are set in a straight line across the front of the premaxillae. 
They are similar in general shape to those of M. monachus, although there is more of 
a " waist " at the junction of root and crown. The canines are similar to those 
of M. monachus. 

The molars are large, the 3rd is largest and the others decrease in size in the order 
3, 2, 4, 1, 5. They are hardly oblique, the 2nd molar being the most so. The molars 
are double rooted except the first, and the last also appears to have only one root in 
the two British Museum specimens. The molars have a low central cusp, and one 
anterior and two posterior smaller cusps. The 5th molar has only a single posterior 
cusp. All the teeth have a well developed cingulum and the crown is slightly rugose. 
Variation and disease. — In the three skulls examined there is no variation or disease 
in upper or lower teeth. 

Lower. — The first lower incisors are smaller than the second, are set internally and 
posteriorly to them and are in a recumbent position. In shape they are similar to 
the upper incisors. The lower canines are similar to the upper ones. 

The 3rd molar is the largest and the others decrease in size in the order 3, 4, 2, 5, 1, 
although molars 3, 4 and 2 are very much of the same size. The teeth are set in the 
line of the jaw, not obliquely. All the molars are double rooted except the first. 
The lower molars are similar in shape to the upper except that the cusps on the 1st 
are rather indistinct, and there is only one small posterior cusp on the 5th. 

The teeth of M. tropicalis examined seem to be more worn than the teeth of 
M. monachus. 

M. schauinslandi 

Dental formula : . 2 1 5 

1 -, c -, m ^ 

2 i' 5 

Upper. — The incisors are set in a straight line across the front of the premaxillae, 
and as far as can be seen from the photographs, the setting and shape of the teeth are 
similar to those of M. tropicalis. The molars are not set obliquely and seem to have 
a main low cusp, a single small anterior cusp and two small posterior cusps. 

Lower. — The lower teeth appear to be similar to those of M. tropicalis. 

Summary of the differences between the teeth 

1. The incisors have a very pronounced " waist " at the junction of root and 
crown in M. tropicalis and M. schauinslandi. 

2. The molars are set very obliquely in M. monachus. 

3. The molars of M. monachus have a large central cusp and single smaller 
anterior and posterior cusps. Those of M. tropicalis and M. schauinslandi have a 
low central cusp, a single small anterior cusp and two small posterior cusps. 



4. The incisors are set straight across the premaxillae in M. tropicalis and M. 
schauinslandi , but on a slight curve in M. monachus. 

Teeth of a very young M. monachus 

There is no exact information about the skull 1892. n. 7.1, 1063I, but from its 
size and the condition of its teeth it is probably new-born. X-rays of the teeth 
were taken (Nos. 764 and 765) (Text-fig. 8). 

5 CMS. 

Fig. 8. — Tracings from X-ray photographs of the teeth of a young M. monachus, Reg. No. 
1892. 1 1. 7. 1. a. Ventral view. b. Oblique view. c. Lower jaw. 


Milk teeth 

These are seen more clearly in the skull itself than in the X-ray. Six milk teeth 
are visible in the upper jaw, three on each side. They are very small, the largest 
being 2 mm. long ; they have no roots and are present in the dried gum over the 
unerupted permanent teeth. The milk molars are placed behind their permanent 
successors. In the upper jaw milk molars 3 and 4 are present on the right side, and 
2 and 3 on the left side. The two outer milk incisors are present. In the lower jaw 
the only milk tooth present is mm 2 on the right side. 

There is no milk predecessor for the first molar in either upper or lower jaws. 

Milk dentition : . 2 1 3 

1 -, c -, m - 

2' i 1 3 

Permanent teeth 

The tips of all the permanent teeth are through the layer of dried skin on the 
skull, though probably apart from the upper canines, none of them would have 
pierced the gum in the living animal. 

The permanent molars are large, but are normal in shape and position. This 
specimen shows an interesting variation in the number of incisors. The upper jaw 
has seven incisors, three of which are supernumerary. On the left side the outer 
incisor is in its normal position, the inner incisor is pushed a little further away from 
the mid line than is normal, and on its median surface, slightly posteriorly is a small 
extra tooth. On the right side the outer incisor is displaced posteriorly by an extra 
tooth which is between it and the canine, and between the inner incisor and the mid 
line is another extra tooth of the same size as the small extra one on the left side. 
The larger extra tooth on the right has the same size and appearance as the inner 

The lower permanent teeth are normal. 

Supernumerary bones in the skull. 

Three of the youngest skulls of M. monachus (1892 .11.7.1, 1063c and 1894 . 7 . 27 . 3) 
have supernumerary bones in the cranium. The youngest specimen (1892. n. 7.1) 
has two bones symmetrically placed in the back of the skull, each bone bounded by 
exoccipital, parietal and supraoccipital. These bones are frequently found in the 
young of other seals and have been known as the tabulare, on the assumption that 
they were homologous with the tabulare of the reptilian skull. Doutt (1942) 
mentions their presence as well as that of two small adjacent bones, in skulls of Phoca 
vitulina and P. hispida and after quoting various authors who have considered the 
relationships of these bones with those in the reptilian skull, he thinks " that it is 
better, for the present at least, to consider these extra bones in the occipital region 
of the seal as being of the nature of fontanelle bones rather than to try to homo- 
logize them with elements in the reptilian skull ". He says that these " extra- 
occipital " bones cannot be considered as Wormian bones for they are too 
symmetrical and too regularly situated. 



Undoubted Wormian bones are also found in this skull and in the two other young 
ones mentioned. In 1892. n. 7.1 they take the form of three small bones, two 
posteriorly and one anteriorly, at the junction of the two parietal bones with the 
frontal bones. In 1063c there is a single triangular bone in the same position. 
Also in the same position in 1894.7.27.3 there are two narrow bones one behind 
the other (Text-fig. 9). 




Fig. 9. — a. "Extra-occipital" bones in M. monachus Reg. No. 1892. 11. 7.1. b. 
Wormian bones in M. monachus Reg. No. 1892. 11. 7.1 ; c. Wormian bones in M. 
monachus Reg. No. 1063c ; d. Wormian bones in M. monachus Reg. No. 1894.7.27.3. 
p., Parietal ; /., frontal ; ex., exoccipital ; soc, supraoccipital. 

Scapula (Text-fig. 10) 

The scapula of Monachus is triangular in shape, the extreme antero-posterior 
length being greater than the height. In this character it differs from the scapulae 
of other Phocids where the anterior edge is much longer and, especially in the 
southern Phocids, much more square in shape. The junction of the posterior and 
dorsal edges is not hook-shaped. The spine is low and poorly developed like those 



of Mirounga and the southern Phocids, and very unlike the plate like spines of 
P. vitulina and H. grypus. The acromion process is well developed and resembles 
that of the southern Phocids, and the glenoid cavity is narrow and kidney-shaped. 
The outer surface of the scapula is convex anteriorly to the spine and concave 
posteriorly to it. Scapulae of P. vitulina and H. grypus are convex posteriorly 
and concave anteriorly. 

10 CMS. 

Fig. io. — Scapulae of a. M. monachus Reg. No. 1894.7.27. 1. 
b. M. tropicalis 1887.8.5. 1. 

In young M . monachus and in M. tropicalis the anterior edge of the scapula sweeps 
round in a continuous curve from the neck, forwards and then back along the dorsal 
edge. In adult M. monachus the anterior edge is more squared off. The scapula 
of M . tropicalis is more elongated antero-posteriorly than that of M. monachus. 

The separate origin of the coracoid is visible in three of the youngest specimens of 
M. monachus. In the youngest scapula (1892. 11. 7.1) it does not appear to have 


begun to ossify. Scapulae 1894.7.27.3, and 1063c have the cartilage still present 
over the glenoid cavity and the coracoid is visible as a distinct bone embedded in it. 
Scapula 1892. 10.4. 1 has no cartilage, the coracoid has fused to the anterior edge of 
the glenoid cavity, but is still recognizable as a separate entity. 


The humerus is short and robust. In common with all the southern Phocids 
there is no supracondylar foramen. This foramen is present in Phoca, H. grypus, 
E. barbatus and C. cristatus. The deltoid ridge is strongly developed and extends 
nearly the whole length of the shaft. The deltoid rugosity on the external surface 
of the ridge is more prominent in M. monachus than in M. tropicalis, but does not 
overhang to so great an extent as in Phoca, Halichoerus and Erignathus. The 
bicipital groove is broad and shallow. In Monachus and the southern Phocids 
there is only a very poorly marked supinator ridge, while this is well developed in 
the northern Phocids. For bones of approximately the same length, the humerus of 
M. tropicalis (PI. 7,a) is slightly more slender than that of M. monachus. The 
distal articular surface is narrower and appears to be slightly more oblique than in 
M. monachus. In M. tropicalis, but not in M. monachus there is a very slight 
anconeal fossa. 

Radius and Ulna. 

The radius is laterally flattened and bowed forward, so that its hinder edge is 
concave. The ulna is compressed laterally, with an expanded olecranon process 
and a concave hinder margin. The ulna of M. monachus is slight y more stoutly 
built than that of M. tropicalis, but presents no special modifications. 

Manus (PL 7,a) 

The carpus consists of seven bones : scapholunar, trapezium, trapezioid, os 
magnus, unciform, cuneiform and pisiform. The length of the digits, and of the 
metacarpals decreases in size from the first to the fifth. The first digit is strongly 
built, the fifth slightly less so, and the second, third and fourth of equal thickness. 
The terminal phalanges are grooved for the insertion of claws. An X-ray taken of 
the manus of an immature M. monachus (1894 . 7 . 27 . 3) shows the presence of distal 
epiphyses on all five metacarpals, but a proximal epiphysis on only the first. All 
the phalanges except the terminal ones have epiphyses on both ends, and a proximal 
epiphysis is visible only on the first terminal phalange, it not yet having ossified 
on the remaining phalanges. 

Pelvis and sacrum 

The innominate bones are of the typical Phocid pattern with short recurved ilia 
and long ischia and pubes. The symphysial area is very small and appears to be 
mainly cartilaginous. The innominate bones of M . monachus have thick flattened 
ilia which are not excavated on their exterior surfaces as in Phoca and Halichoerus, 



When compared with the innominate bones of other Phocids it is seen that the ilia 
of M. monachus are everted and also are rotated upwards relative to the plane of 
the pubes and ischia, so that the external surfaces of the ilia face slightly posteriorly 
and dorsally. In Phoca and Halichoerus and also in the southern Phocids the ilia 
are everted in approximately the same plane as the pubes and ischia. The ischium 
of M . monachus is strongly built, approximately circular in cross section until about 
the level of the ischiatic spine, when it continues as a flat bar. The pubis is also 
thicker at its origin and flattens out posteriorly to meet the ischium. The ischium 
forms the upper and practically the whole of the posterior border of the obturator 
foramen before it meets the pubis. 

A comparision of the innominate bones of Phocids shows that in the northern 
genera Phoca, Halichoerus and Erignathus the obturator foramen is long and narrow 
and the width of the innominate at the level of the ischiatic spine less ; in the 
southern genera Ommatophoca, Hydrurga, Leptonychotes, Lobodon and Mirounga the 
obturator foramen is broad and the width of the innominate correspondingly 
greater. The proportions of the obturator foramen given in the table below indicate 
a closer relationship of Monachus with the Southern than the Northern Phocids 
(PL 8). 

Measurements of the Obturator Foramen 







of width 



to length. 

P. vitulina .... 96 

3 1 


H. grypus 




E. barbatus . 




L. weddelli 




H. leptonyx . 




L. carcinophagus 




0. rossi 




M. monachus 

io63g 94 



io63f 98 



1421a 82 


60 9 

M. tropicalis 

1887.8.5. 1 ... 112 







In M . monachus the pectineal tubercle and the ischiatic spine are well developed, 
the acetabulum is deep and circular with a well marked cotyloid notch. The 
innominate bones articulate with two sacral vertebrae, but in the three adult M. 
monachus, two have four vertebrae fused to form the sacrum, and one — the 
youngest — has three. The wings of the sacrum are broad and strong, and 
articulate on their posterior surfaces with the ilia. 


In most respects the pelvis of M. tropicalis is similar to that of M. monachus, but 
the size of the ischium and pubis differs considerably. The pubis is extremely 
strongly built, is circular in cross section, tapers slightly towards its posterior end 
and does not flatten out except at the symphysial area. The ischium is much more 
slender than the pubis, and posterior to the ischiatic spine it is only a very narrow 
bar, triangular in cross section. 


The femur is short and flattened antero-posteriorly. The greater trochanter is 
large and thickened and is separated from the head by a groove which is deeper 
and more distinct in M. monachus than in M. tropicalis. In common with all Phocids 
except Phoca and Halichoerus there is no trace of a digital fossa. The distal end of 
the femur is very broad because of the great development of the epicondyles. In 
general shape, and in particular in the great width in proportion to its length, the 
femur of Monachus is more like that of the Southern than the Northern Phocids. 


The patella of M. monachus is a small flat bone articulating with the femur. The 
measurements of specimen i033f are : Greatest antero-posterior length 26 mm. 
Greatest height anteriorly 15 mm. The patella of M. tropicalis, while of the same 
general length, is, particularly in the Cambridge specimen, higher and more 
pyramidal. The measurements of the British Museum and Cambridge specimens 
respectively are : Greatest antero-posterior length 30 mm., 24 mm. Greatest 
height anteriorly 20 mm., 28 mm. 

Tibia and Fibula 

The tibia and fibula are of the usual pinniped form, the two bones being fused at 
the proximal end, but separate from each other at the distal end. In only one of 
the M . monachus in the collection (io63g) is one of the fibulae fused to the tibia, so 
this fusion must be one of the last to occur. The posterior tibial fossa is less concave 
than in any of the other Phocids examined, but the tibial spine and the anterior 
tibial fossa are quite well marked. Both the tibia and the fibula articulate at their 
lower ends with the astragalus. 

Pes (PI. 7,b) 

The tarsus consists of seven bones : astragalus, calcaneum, navicular, cuboid and 
external, middle and internal cuneiform bones. The length of the digits and of 
the metacarpals decrease in size in the order 1, 5, 2, 4, 3, the middle digit being the 
shortest. The outer digits are the most strongly built. The terminal phalanges 
are grooved for the insertion of small claws. An X-ray taken of the pes of a young 
M. monachus (1894.7.27.3) shows the presence of distal epiphyses on all the 
metatarsals, and a proximal one on the first only. Epiphyses are visible on both 



ends of all the phalanges except the terminal ones. Epiphyses have not yet ossified 
on the proximal ends of any of the terminal phalanges except the first. 

Vertebral Column 

The vertebral formula is as follows : 

M . monachus 






1892. 1 1. 7. i, 1063I. 





3 + 

1894.7.23.3, 1063I1. . 






1894.7.27.2, io63g. 






1894.7.27. 1, io63f 






1892. 10.4. 1, 1063d 






1863. 4. 1. 1, 1421a 












M. tropicalis 

1887.8.5. 1 






899c . 






Cervical vertebrae 

M. monachus. — The centra are approximately circular in cross section and the 
ventral surfaces have a median keel in all except the atlas. The neural arches are 
narrow antero-posteriorly, the widest (the 7th) being 20 mm., and the neural spines 
increase from a hardly perceptible point on the third vertebra to a spine 38 mm. 
high on the seventh. The transverse processes are perforated by the vertebrarterial 
canal in all except the seventh. The transverse processes of cervical vertebrae 3-6 
inclusive are, in all the Phocidae examined, divided into two branches, a dorsal 
transverse element which is more or less at right angles to the median plane, and 
directed slightly posteriorly, and a ventral costal element which is directed more or 
less vertically downwards and is expanded antero-posteriorly into a plate which is 
greatest in the sixth vertebra. In M . monachus the transverse processes leave the 
centrum at an angle of approximately 45 ° and are not divided into two branches 
(Text-fig. 11). 

The cervical vertebrae of M. tropicalis, while conforming to the same general 
pattern as those of M. monachus, are different in several minor respects. They 
give the general impression of being more finely built than those of M. monachus. 
The transverse process of the atlas is not so massive and the vertebrarterial canal is 
much larger, the neural spine of the axis is not so high and does not project so far 
backwards. The transverse process of the axis is thin and pointed, those of verte- 
brae 3-6 are divided into two branches, the costal element not being so expanded 
as in Phoca and directed not vertically, but laterally at an angle of approximately 
45°, and inclining posteriorly. The general shape of the cervical vertebrae and in 
particular that of the transverse process is more like Leptonychotes than Phoca. 

It is interesting to note that both in the British Museum and the Cambridge 
specimens of M . tropicalis the neural arches of the third and fourth vertebrae do 



not meet dorsally. In the British Museum specimen the two sides of the neural 
arch of the fifth vertebra are fused, but the lateral tips of the spine are curved 
outwards, to give a bifurcated tip to the spine. The lateral tips of the sixth spine 
are less curved. In the Cambridge specimen the two sides of the fifth neural arch 

Fig. 11. — a. 5th cervical vertebra of M. monachus Reg. No. 1894.7.27.2. b. 5th 
cervical vertebra of M. tropicalis Reg. No. 1887.8.5. 1. c. 2nd lumbar vertebra of 
M. monachus Reg. No. 1894.7.27.2. d. 2nd lumbar vertebra of M. tropicalis Reg. 
No. 1887.8.5. 1. 

meet, but are not fused, and the tips curve outwards. The spine of the sixth is 
fused with curved tips, and the spine of the seventh in both animals is normal. A 
similar non-fusion of the neural arch occurs in M. monachus 1063c, where the two 
sides of the third neural arch fail to meet. The remaining arches are normal and 
do not have the bifurcated appearance of those of M. tropicalis. Ward (1887b) 



says, describing M. tropicalis, "At no time does this seal raise its head as much 
above the line of its back as does the harbor seal : the flexibility of its cervical 
vertebrae appearing to be quite restricted ". It is difficult to know whether this 
is a character that applies to M. tropicalis in general, or only to the animals that 
Ward saw ; or whether it has any connection with the non-fusion of the neural 
spines, as the two animals in which this character is present were both collected by 
Ward from a single group of animals. 

Thoracic vertebrae 

The thoracic vertebrae of M. monachus articulate with fifteen pairs of ribs. Ribs 
i and 2, and n, 12, 13, 14 and 15 articulate only with the corresponding thoracic 
vertebra, but the capitular articulation of ribs 3-10 inclusive overlaps on to the 
centrum of the vertebra in front. Ribs 1-13 inclusive articulate by both capitular 
and tubercular heads, the two articulations getting closer together until ribs 14 and 
15 articulate by means of a single head only. The transverse processes are 
prominent, but diminish in size from the first to the tenth vertebra and are hardly 
visible from the eleventh to the fifteenth. The neural spines are high and pointed 
in the first five vertebrae, but get progressively smaller and more backwardly 

Between thoracic vertebrae 9 and 10 in specimen 1863 .4. 1 . 1, the anterior common 
ligament has ossified in the form of a bony plate, which is fused to the centrum of 
the tenth vertebra and extends anteriorly for 4cm. beyond it. Evidences of a 
similar abnormality, though to a much lesser extent, are present on the ventral 
surfaces of the centra of vertebrae n, 12 and 13. 

The thoracic vertebrae of M. tropicalis are practically the same as those of M. 
monachus except that the neural arches are wider and the neural spines are shorter 
and less pointed. 

Measurements of thoracic vertebrae 

Length medially of neural arch of 10th vertebra 
Least width ,, ,, 

Height* of neural spine of 1st vertebra 

Length of centrum of 1st „ 

„ „ „ „ 10th 

M. monachus. M. tropicalis. 


1887.8.5. 1. 

















Height not taken vertically, but medially along spine. 

Lumbar vertebrae 

The lumbar vertebrae of M. monachus have large heavy centra that are slightly 
concave ventrally. The transverse processes are prominent and project anteriorly 
and ventrally. The cephalic articular processes are large and are directed obliquely 



anteriorly, but the caudal articular processes are extremely small and thin, and may 
not even reach the vertebra behind. The neural spines are stoutly built and are 
directed slightly posteriorly. 

The right transverse process of the first lumbar vertebra of M. monachus io63f 
is normal, but the left is stout, truncated, and its distal end forms an articulation 
for a small triangular " pleurapophysial " ossicle 34 mm. in length. 

A condition of chronic osteo-arthritis is present on the posterior half of the centrum 
of the fifth lumbar vertebra of M. monachus io63g, and this condition has spread to 
the centrum of the first sacral vertebra. 

The lumbar vertebrae of M. tropicalis are in general like those of M. monachus, 
though the articulations are normal, the caudal articular process overlapping to a 
considerable extent the cephalic articular surface of the next posterior vertebra. 
The neural spines are laterally flattened, low, rounded, and either vertical or 
inclined slightly anteriorly (Text-fig. 11). 

Caudal vertebrae 

Except for the first two or three, the caudal vertebrae are without a neural arch, 
and both the number and size of the processes, and also the size of the centrum, 
diminish in size from before backwards. 


The articulations of the ribs with the thoracic vertebrae have already been 
discussed. In M. monachus (io63f) the ribs increase in length from the 1st (yy mm.) 
to the nth (263 mm.) and then decrease to the 15th (209 mm.). In M. tropicalis 
(1887. 8. 5.1) similar measurements are 1st (71 mm.), nth (255 mm.) and 15th 
(189 mm.). 

The adult specimens of M. monachus in the collection lack the cartilaginous 
portions of the ribs. An immature M. monachus (1063c) has cartilaginous ribs 
attached directly to the sternum from bony ribs 1-9 inclusive. Cartilaginous ribs 
10-15 inclusive turn forward and lie against the cartilaginous portion of the rib 
in front. 

Cartilaginous ribs 1-10 inclusive of M. tropicalis (1887. 8. 5.1) are attached 
directly to the sternum, 11 and 12 are long and lie against the cartilaginous portion 
of the ribs in front, but the cartilaginous portions of ribs 13-15 are very short, 
between 35 and 70 mm. long, and are unattached ventrally. 

The first cartilaginous rib articulates with the manubrium of the sternum, the 
remaining nine with the cartilages between the sternebrae, ribs both 8 and 9 
articulating with the last cartilage. 


The sternum of both M. monachus and M. tropicalis consists of nine sternebrae, 
although there are only eight present in M. monachus 1063c. The sternebrae are 
dorso-ventrally flattened and more or less quadrate in shape, the first and last being 



more elongated. The xiphisternum is prolonged posteriorly into two cartilaginous 
extensions, the ends of which are expanded and joined together posteriorly. 


In view of the small number of specimens available no attempt at age determina- 
tion has been made but it has been possible to do a certain amount on the growth 
of the skull and skeleton. 

Following Doutt's (1942) method for estimating the " suture age " of the skull 
the following table was drawn up. (Table II). As suture closure is a gradual 
process the degree of closure has been given a value : 1 for open, 2 for less than 
half closed, 3 for more than half closed, and 4 for completely closed, and the total 
for each skull is known as the " suture age " for that specimen. 

Table II. — Suture Ages of the Skulls 















































h- 1 











•— > 


















M. monachus : 

1892. 11. 7. 1 









. 171 










. 217 

1894.7.27.3 <J 









. 224 

1892. 10.4. 1 









. 260 










. 262 

1894.7.27.2 $ 











. 268 

1863. 4. 1. 1 £ 











• 273 

1894.7.27. 1 <J 











. 281 











29 + 

• 295 

M. tropicalis : 

1889. 11. 5. 1 














. 267 

1887.8.5. 1 

The ore 


.er of 





are is 


as f ol 


lows : 




• 3 




. 269 




— bas 





ito — ] 









Basioccipito — basisphenoid. 
7. Basisphenoid — presphenoid. 

In Table II the skulls are placed in order of condylo-basal length, and it can be 
seen that the one known female skull of M. monachus has a higher " suture age " in 



proportion to its condylo-basal length than the other skulls, i.e. it matures at a 
smaller size. For this reason it seems likely that the skull and skeleton of M. 
tropicalis 1887.8.5. 1 are also from a female. 

Table II also shows that the series of skulls of M. monachus, although more or 
less uniformly distributed over a size range, is divisible by " suture age " into two 
discontinuous groups — those with " suture ages " 12-18 and those from 26-30. 

This is clearly shown in the table of percentage increases below, from which it is 
also evident that while the " suture age " increases from 12-18, an increase of six 
units, there is a general increase in the size of all the components of the skull, the 
average amount being 53% or 9% per unit of " age ". During the period when the 
" suture age " increases by four units from 26 to 30 there is also an increase in all 
the components with one exception, but the average rate of increase per unit of 
" age " is now only 4.5%. A diminution in the rate of increase with age is to be 
expected, but one component — the cranium — appears virtually to have ceased 
growing by the time a " suture age " of 18 is reached ; there are no crania in the 
26-30 group larger than the largest individuals in the 12-18 group. 




Percentage increases. 










Snout Width at 







length. canines. 







• 59 • 39 


. 44 



. Negative 


8 . Negative . 







. 24 . 25 


• 15 

The most striking feature of the above table is that, despite the absence of any 
large size discontinuity in the series, there is a complete absence of any specimen in 
the large " suture " group 19-25 ; the largest specimens of the " younger " 12-18 
group are almost as large as, and in some components larger than the smallest 
individuals of the " older " 26-30 group. 

It is not likely that growth proceeds in this erratic fashion and therefore it is 
concluded that the " suture age " is not a rectilinear age index. It seems clear 
that at a certain stage of growth there is a very rapid suture closure. After this 
stage is past the skull components continue growing with the notable exception of 
the cranium. Skulls with a condylo-basal length of less than ± 265 mm. have a 
rapidly growing cranium and during this growth-phase the occipito-parietal, 
sagittal, coronal and basioccipito-basisphenoid sutures are closing. At ± 265 mm. 
condylo-basal length there is rapid suture closure affecting the occipito-parietal, 
squamoso-parietal, sagittal, coronal, basioccipito-basisphenoid and basisphenoid- 
presphenoid sutures, and cranial growth ceases. After this, growth of the facial 
elements continues and the interfrontal, intermaxillary, fronto-maxilla, jugal- 
maxilla, squamosal- jugal and palato-maxilla sutures are open in all the skulls of 
M. monachus in the collection. 

It is possible that the " suture age " figures give an approximation to a rectilinear 
age index if the 19-25 period is eliminated, and if this is done the relative " ages " 



of the skulls are 12 : 12 : 13 : 16 : 18 : 23 : 19 : 21 : 23. The " suture age " 12 
includes two skulls which are both young but which are clearly at very different 
stages of growth. The youngest skull — 1892. n. 7.1 — has not been included in the 
graph as it is so much younger than its " suture age " indicates, this criterion being 
invalid for very young (and probably very old) skulls. 

Plotting the skull measurements against this time scale (Text-fig. 12) gives a 




IR 13 14 15 Ifc 17 18 19 20 21 XX Z7> 

Suture age 
Fig. 12. — Graph showing relative grtfwth rates of different regions of the skull. 



picture which is not an unreasonable one, and one which suggests certain sex 
differences. Even if the time scale is only an approximation, plotting all the bones 
to this scale gives a series of curves that are directly comparable. The slope and 
shape of these curves show the relative growth rates and growth patterns of the 
different elements. 

From a visual inspection of a series of skulls of M. monachus of ascending size, it 
is obvious that the various regions of the skull do not all grow at the same rate. 
The cranium for instance, while it increases in length, does not do so nearly so quickly 
as the facial region. Measurements of the cranial, interorbital and snout regions 
were taken between uprights and these, together with width measurements are seen 
in Table III. The interorbital region is measured to the maxillary tubercle at the 
anterior edge of the orbit. 

The logarithmic values of the various measurements were plotted in turn against 
the logarithmic value of the condylo-basal length. The points fall approximately 
about a straight line, showing that there is allometric growth. This being so, it 
may be taken that the rates of growth of the various parts of the skull relative to 
the condylo-basal length satisfy the equation y=bx fc , where x is the condylo-basal 
length, y the length of the part, b the fractional coefficient (the value of y when 
x=i) and k the growth coefficient. 

Registered No. 

M. monachus 
1892. 11 .7.1 


10.4. 1 



4. 1.1 





Table III 

Condylo- Inter- 
basal Cranium orbital Snout 
length. length. length. length, 
mm. mm. mm. mm. 

171 . 85 

217 . 104 

224 . 101 

260 . 124 

262 . 118 

268 . 112 

273 . 114 

281 . 113 

295 . 117 


width at 




. 40 




. 58 



137 • 












159 • 








185 • 

. 89 




. 87 




Width at 
upper edge 


The growth coefficient is the tangent of the angle between the horizontal and the 
line joining the points. Values of k greater than unity indicate an increasing rate 
of relative growth, and those less than unity the converse. The following are the 
values of k for the various parts : 

Cranium length . . . .0-62 

Interorbital length 

. i-8i 

Snout length 

. 1-32 

Snout width 

• 0-95 

Zygomatic width . 

• I'I5 

Auditory meatus width 

. 0-85 

ZOOL. 3, 5. 



These figures bear out the visual evidence as they show that the interorbital and 
snout regions, and the zygomatic width have a much faster relative growth than 
the rest of the skull, while the cranium, the snout width and the width of the cranium 
between the external auditory meatuses grow at a much slower rate. 

Although fusion of the bones of the skull commences at an early age, the epiphyses 
of the skeleton do not fuse until comparatively late. The following is the order of 
fusion of the epiphyses and skeletal elements : 

i. Pelvic elements. 
2. Sacral vertebrae. 

Humerus (distal). 

Femur (proximal). 
5. Vertebral epiphyses. 

Humerus (proximal). 

Radius (proximal). 

Femur (distal). 

Tibia (proximal). 

Fibula (proximal). 
11. Ulna (proximal). 

Metacarpal epiphyses. 

Metatarsal epiphyses. 

14. Fibula to tibia. 

15. Radius (distal). "^ 

Ulna (distal). I Not fused in any specimen 

Tibia (distal). f in collection. 

Fibula (both epiphyses). J 

The above list shows that the epiphyses of the limb bones tend to fuse first to 
the bones nearest the body — humerus and femur — and the fusion proceeds outwards 
to the digits. In the youngest animal (1892. 11. 7.1) the separate elements of the 
vertebrae have not yet fused, and the fusion of the vertebral epiphyses to the centra 
in the older animals appears to start at the cervical end of the column and proceed 

In order to see how the limb bones and pelvis increase in length with age, 
measurements were taken of the lengths of the pelvis and of the shafts of five limb 
bones, not including the epiphyses, of three male animals of increasing age, and the 
new-born specimen, as there can be no difference due to sex at this age. There are, 
unavoidably, many inaccuracies in the table (Table IV). The shaft length is 
difficult to measure exactly in the two youngest animals because of the large amount 
of cartilage present where the epiphyses are not fully ossified. The proportions, 
using the measurements of the largest specimens as 100%, are taken on the assump- 
tion that all the animals will grow to this size. This is obviously not so, and even 
this animal is not fully mature, but it is the oldest that is known to be male. The 
table shows that the pelvis has over half (62.2%) of its growth in length to complete 
after the animal is born. The corresponding figures for the limb bones are : 
humerus 45.4% radius 50%, femur 54.3%, and tibia 55-4%- 


Table IV 








mm. % 

mm. % 

mm. % 

mm. % 

mm. % 

M . monachus : 

1892. 11. 7. 1 

59 54'6 

63 50-0 

• 43 45'7 

91 44-6 

92 37-8 

1894.7.27.3 c? . 

76 70-0 

80 65-1 

■ 58 6i-7 

127 62'2 

147 60-4 

1863. 4. 1. 1 6" • 

101 93*5 

. 118 93-6 

. 88 93-6 

l86 91- I 

221 90-9 

1894.7.27. 1 6*. 

108 100 

126 100 

94 100 

204 IOO 

. 243 IOO 


Family Phocidae Gray, 1825. 
Subfamily Monachinae Trouessart, 1904. 

The Phocidae are divided into the subfamilies Phocinae, including Phoca, Erig- 
nathus and Halichoerus, with the incisive formula 3 ; the Cystophorinae including 

Cystophora and Mirounga, with incisors 2 ; the Lobodontinae, including Lobodon, 

Hydrurga, Leptonychotes, and Ommatophoca with incisors 2 ; and the Monachinae 

with the single genus Monachus also with incisors 2. 


The Cystophorinae are quite distinct, and are not considered here. Some of the 
characters which distinguish the Phocinae from the Lobodontinae are : the 
zygomatic process of the maxilla with the posterior border subvertical, not extending 
far backwards beneath the malar ; nails of all the digits well developed ; and the 
outer digits of the pes not much prolonged beyond the others. In the Lobodontinae 
the zygomatic process of the maxilla is prolonged backwards beneath the malar ; 
the nails of the hind limbs rudimentary ; and the outer digits lengthened. 

A comparison of members of the genus Monachus with members of the two 
preceding subfamilies shows that in the skull the incisive formula is the same as in 
the Lobodontinae, the extension backwards of the zygomatic process of the maxilla 
is not quite so great as in the latter subfamily, but is very much greater than in the 
Phocinae. All members of the Phocinae have large claws on both fore and hind 
flippers, while Monachus agrees with the Lobodontinae in having the hind claws 
very much reduced. A further similarity is in the shape of the hind flippers which 
have the two outer digits very much longer than the inner ones, the first digit of the 
fore flippers is the longest and the rest get gradually shorter. In the Phocinae the 
fore flipper is much more square in shape and the digits are more nearly equal. 
The Phocinae also differ in the position of their nostrils, which are on the anterior 
end of the snout and more or less vertical. The nostrils of both the Monachinae 
and the Lobodontinae are nearly on the dorsal surface of the snout and are almost 
horizontal. As already noted in the various sections on the bones of the skeleton 
of Monachus, these are in general much more like those of the Southern than the 


Northern Phocids. The above evidence shows clearly that the Monachinae are 
more closely related to the Lobodontinae than to the Phocinae. 


I have to thank Dr. G. von Wahlert of the Ubersee-Museum, Bremen, and Dr. 
K. Zimmermann, Zoologisches Museum der Universitat, Berlin, for help in locating 
the type specimen of Monachus schauinslandi and for information and photographs 
relating to it ; and also Dr. F. R. Parrington of the University Museum of Zoology, 
Cambridge for access to an articulated skeleton of M. tropicalis. I am very much 
indebted to Dr. H. W. Parker, Keeper of Zoology, for his advice and particularly 
for his assistance with the section relating to the growth of the skull. I am also 
very grateful to Miss Hilary King and to Commander J. M. Chaplin, R.N. (Ret.) for 
much help in the translation of papers. 


Anon. 1894. List of additions to the Society's Menagerie during the year 1894. Proc. Zool. 
Soc. Lond. 1894 : 749. 

1903. The West Indian Seal. Bull. N.Y. Zool. Soc. No. 9 : 83. 

1910a. Additions to the Society's Menagerie. Proc. Zool. Soc. Lond. 1910 : 768. 

19106. Rare Tropical Seals. Bull. N.Y. Zool. Soc. No. 38 : 644. 

1926. Mediterranean Seals for the Museum. Nat. Hist. N.Y. 26 : 656. 

1951. Hawaiian Monk Seal, marine rarity. Zoonooz 24 (7) : 3. 

— ■ — 1954. Failure of grass to check erosion. Times, Friday, nth June, 1954. 
Agacino, E. M. 1945a. Algunos datos sobre cierto mamiferos del Sahara Occidental. Bol. 
Soc. Esp. Hist. Nat. 43 : 199-212, 2 pis. 

19456. Las Focas del Sahara. Cal. Mens. Illustr. de Caza y Pesca. Armas y Guarderia, 

26 : 45-46, 1 fig. (not seen). 

1945c. Las Focas del Sahara. A.O.E. 3 : 64-65, 2 figs. Sidi Ifni (not seen). 

1950. Notes sur les Phoques-Moines [Monachus monachus Herm.) du littoral Saharien 

Espagnol. Mammalia, 14 : 1-6, pi. i and ii, fig. 1. 

Aharoni, J. 1930. Die Saugetiere Palastinas. Z. Sdugetierk. 5 : 327-343. 

Allen, G. M. 1942. Extinct and vanishing mammals of the Western Hemisphere. Spec. 

Publ. Amer. Comm. Int. Wildlife Prot. 11 : 1-620, figs. 
Allen, J. A. 1880. History of North American Pinnipeds. U.S. Dept. Int. Geol. and Geogr. 

Survey. Misc. Publ. 12, xvi + 785, figs. 
■ 1887a. The West Indian Seal [Monachus tropicalis Gray). Bull. Amer. Mus. Nat. Hist. 

2 : 1-34, pis. i-iv. 
— — 18876. The West Indian Seal. Science, 9 : No. 206, 35. 

1887c. The West Indian Seal. Science 9 : No. 207, 59. 

— — 191 8. The Laysan Seal. Nat. Hist. N.Y. 18 : 399-400. 

Alessandrini, A. 1819a. Osservazione su gl'inviluppi del feto della Phoca bicolor. Opuscoli 
scientifici. 3 : 298-316. 

18196. Bemerkungen iiber die Fotushiillen der Phoca bicolor. Arch. Physiol. 5 : 604-616. 

Anderson, J. and De Winton, W. E. 1902. Zoology of Egypt. Mammalia. London, 

xvii + 374, 63 pis., map. 
Antipa, G. 1916. Pescaria si Pescuitul in Romania. Publ. Fond. V. Adamachi 8 : No. 46 

1941. Marea Neagra. Vol. 1. Publ. Fond. V. Adamachi 55 : 1-3 13. 

Atkinson, A. L. C, and Bryan, W. A. 1913. A rare seal. Bull. N.Y. Zool. Soc. 16 : 1050- 


Bailey, A. M. 1918. The Monk seal of the Southern Pacific. Nat. Hist. N.Y. 18 : 396-399. 

1952. The Hawaiian Monk seal. Mus. Pictorial Denver, 7 : 1-32. 

Barcelo y Combis, F. 1875. Apuntes para la fauna Balear. An. Soc. Esp. Hist. Nat. 

4 : 53-68. 
Baylis, H. A. 1937. O n the Ascarids parasitic in seals with special reference to the genus 

Contracoecum. Parasitology. 29 : 1 21-130. 
Bertram, G. C. L. 1943. Notes on the present status of the Monk Seal in Palestine. /. Soc. 

Pres. Fauna Emp. No. 47 : 20-21. 
Blackman, T. M. 1941. Rarest seal. Nat. Hist. N.Y. 47 : 138-139. 

Blasius, J. M. 1857. Fauna der Wirbelthiere Deutschlands ... 1 Sdugethiere. Braun- 
schweig, pp. vi + 549. 
Bobrinskii, N. A. 1944. iV - The Order Pinnipeds. In Bobrinskii, N. A., Kuznetzov, B. A., 

Kuzakin, A. P. Key to Mammals of the U.S.S.R. Moscow, pp. 1-440, 256 figs., 32 pis., 

62 maps. 
Boddaert, P. 1785. Elenchus Animalium. 1 : pp. xxxviii + 174. Rotterdam. 
Boettger, C. R. 195 1. Notizen zur Verbreitung und fiber die Verwandtschaftsbeziehungen 

der Monchsrobbe (Monachus albiventer Bod.). Zool. Anz. 147 : 303-310. 
Boetticher, H. v. 1930. Die elemente der bulgar Saugetierfauna und ihre geograph, und 

okologischen Grundlagen. Mitt. Naturwiss. Inst. Sofia 6 : 33-42. 
Borcea, I. 1926-27. Donnees sommaires sur la faune de la Mer Noire. Ann. Sci. Univ. 

J assy, 14:536-581. 
Brusina, S. 1889. Siazvci Jadramskogamora. Rad. Jugoslav. Akad. Zagreb, 95 : 79-177. 
Bryan, W. A. 1915. Natural History of Hawaii. Pp. 596, 117 pis. Honolulu. 
Budker, P. 1945. Pinnipedes et Sireniens d'Afrique. Notes Afr. nr. 27 : 4-6. 
Buffon, G. L. de. 1782. Histoire Naturelle. Supplement VI. Paris. 

Cabrera y Latorre, A. 1914. Fauna Iberica, Mamiferos. Madrid, pp. xviii + 441, 22 pis. 
• 1932. Los Mamiferos de Marruecos. Trab. Mus. Cienc. Nat. Madr. Zool. no. 57 : 1-361, 

34 figs., 12 pis. 
Cadogan, F. 1945. Monk seals at Madeira. /. Soc. Pres. Fauna Emp. 52 : 54-55. 
Calinescu, R. I. 1936. Focele dela Capul Caliacra. Bui. Soc. Nat. Roman. No. 8 : 6-10. 
Carrucio, A. 1893. Su di un Pelagius monachus $ adul. e del suo Feto presi a Cap Teulada 

nel Mediterraneo. Boll. Soc. Romana Zool. 2 : 201-21 1. 
Carus, J. V. 1893. Prodromus Faunae Mediterraneae, 2 : 718. Stuttgart. 
Caspers, H. 1950. Beobachtungen fiber das Vorkommen der Monchsrobbe {Monachus 

albiventer Bodd.) in Schwarzen Meer. Neue Ergebnisse und Probleme der Zoologie 

{KlattF estschrift) . Leipzig, pp. 91-105. 
Chapin, E. A. 1925. Descriptions of new internal parasites. Proc. U.S. Nat. Mus. 68 : 

Art. 2, 1-4. 
Clark, J. G. D. 1946. Seal hunting in the Stone Age of North Western Europe : A study in 
economic prehistory. Proc. Prehistoric Soc. 1946 : 12-48. 

■ 1952. Prehistoric Europe. The Economic Basis. London, pp. xvii + 349, 16 pis. 

Cousteau, J. Y., and Dumas, F. 1953. The Silent World. London, pp. xi + 148, illust. 
Cuni, L. A. 191 8. Contribucion al estudio de mamiferos acuaticos obervados en las costas de 

Cuba. Mem. Soc. Cubana. Hist. Nat. 3 : 83-123. 
Cuvier, F. 1 8 13. Description Zoologique d'un Phoque Moine femelle. Ann. Mus. Hist. 

Nat. Paris, 20 : 387-392. 
1824. De quelques especes de Phoques et des groupes generiques entre lesquels elles se 

partegent. Mem. Mus. Hist. Nat. Paris, 11 : 174-214, pis. 12-15. 

■ 1826. Dictionnaire des Sciences Natur elles, 39 : 1-559. Paris. 

Cuvier, G. 1823. Recherches sur les Ossemens Fossiles. Paris, 2nd ed. 5 : pt. 1, illustr. 

D'albertis, 1877-78. Crociera del Violante. Ann. Mus. Storia nat. Genova, 290-324. 

Dampier, W. 1705. Voyages and Descriptions. London, 3rd ed. II : part II, 1-132. 
Dathe, H. 1934. Ein Beitrag zur Wirbeltierfauna Dalmatiens. Zool. Gart. Leipzig, n.f., 

7 : 108-130. 



Dieuzeide, R. 1927. Sur quelques points d'anatomie du Phoque-Moine de la Mediterranee 

(Monachus albiventer Bodd.). Bull. Sta. Aquic. Peche Castiglione, 2 : 215-249, figs. 
Doutt, J. K. 1942. A Review of the Genus Phoca. Ann. Carneg. Mus. 29 : 61-125. 
Du Tertre, J. B. 1667. Hist. Gen. des Antilles. Paris, 2 : 539 pp. 
Eichwald, E. 1841. Fauna Caspio-Caucasia. Nouv. Mem. Soc. Nat. Moscou, 7 : 34. 
Elliot, D. G. 1904. The land and sea mammals of Middle America and the West Indies. 

Field Mus. Publ. Zool. 4 : 441-850, pis. 42-68. 
Elliott, H. W. 1884. The Monk seal of the West Indies, Monachus tropicalis Gray. Science 

3 : No. 72, 752-753- 
Elschner, C. 1915. The Leeward Islands of the Hawaiian Group. Honolulu, pp. 1-68, text 

Fleming, J. 1822. The Philosophy of Zoology. Edinburgh and London, 2 : 1-618, illust. 
Flower, S. S. 1932. Notes on the Recent mammals of Egypt, with a list of the species 

recorded from that kingdom. Proc. Zool. Soc. Lond. 1932 : 369-450. 
Flower, W. H. 1882. Additions to the Society's Menagerie. Proc. Zool. Soc. Lond. 1882 : 

Galvagni, E. 1902. Beitrage zur Kenntnis der Fauna einiger dalmatinischer Inseln. 

Verh. Zool.-Bot. Ges. Wien. 52 : 362-369. 
Gaumer, G. F. 1917. Monografia de los Mamiferos de Yucatan. Mexico, pp. xli + 331 : 

58 pis. 
Gavard, 1927. Observations sur le phoque moine Monachus albiventer Bodd. faites au 

laboratoire de Castiglione. Bull. Sta. Aquic. Peche Castiglione, 2 : 175-21 1. figs, and 

Gervais, F. L. P. 

Atlas, 84 pis. 
Giebel, C. G. A. 

GlSTEL, J. 1848. 

32 p;. 
Gloger, C. W. L. 

1859. Zoologie et Paleontologie Francaises. Paris, 2nd ed., pp. viii + 544, 

1848. Ersch v. Gruber. Allgemeine Encyclopaedic 24 : 284-292. 
Naturgeschichte des Thierreichs fur hohere Schulen. Stuttgart, pp. xvi 



Smithson Misc. Coll. 115 

1841. Gemeinnutziges Hand- und Hilfsbuch der Naturgeschichte. 

1 : pp. xliv + 495. (Seals, p. 160-164). 
Goldman, E. A. 195 1. Biological investigations in Mexico. 

xiii -f 476 pp., 70 pis. 
Gosse, P. H. 1 85 1. A Naturalist's Sojourn in Jamaica. London, pp. xxiv + 497, pi- vn - 
Gray, J. E. 1837. Description of some new or little known mammalia principally in the 

British Museum Collection. Mag. Nat. Hist., n.s. 1 : 577-587. 
— ■ — 1850. Catalogue of the specimens of Mammalia in the collection of the British Museum. 

London, Pt. 2, Seals, pp. v -f 48, 16 figs. 
— ■ — 1854. Description of a new genus and species of seal (Heliophoca atlantica) from Madeira. 

Ann. Mag. Nat. Hist. (2) 13 : 200-202. 
■ 1864. Notes on seals (Phocidae), including the description of a new seal (Halicyon 

richardii) from the west coast of North America. Proc. Zool. Soc. Lond. 1864 : 27-34. 
■ 1866. Catalogue of Seals and Whales in the British Museum. London, 2nd ed., pp. 

vii + 4°2. 101 figs. 
Gruvel, M. A. 1924. Quelques observations zoologiques faites au cours d'un voyage en 

Mauritanie. Bull. Soc. Nat. Acclim. France, 71 : 13-14. 
Gunter, G. 1947. Sight records of the West Indian seal, Monachus tropicalis (Gray), from the 

Texas coast. /. Mammal. 28 : No. 3, 289-290. 
Hayes, J. G. 1928. Antarctica. London, pp. xv + 448, 16 pis., 14 figs. 
Heim de Balsac, H. 1936. Biog6ographie des Mammiferes et des Oiseaux de l'Afrique du 

Nord. Bull. Biol. Suppl. 21 : 1-446, pis. i-vii, maps. 
Hermann, J. 1779. Beschreibung der Miinschs-Robbe. Beschdf. Berlin Ges. Naturf 

Freunde, 4 : 456-509, 2 pi. 
Hill, R. 1843. The Jamaica Almanack for 1843. 


Hill, R. 1846. Description of a seal found on a shoal south of Jamaica. Proc. Zool. Soc. 

Lond. 1846 : 80-81. 
Joyeux, Ch., and Baer, J. G. 1936. Faune de France, 30 : Cestodes. 613 pp. 
Keller, O. 1887. Thieve des Classischer Alterthums. Innsbruck, pp. ix + 488. 

1909. Die Antike Tierwelt. Leipzig, 1 : pp. xii + 434. 

Kellogg, Remington. 1936. A Review of the Archaeoceti. Publ. Carnegie Instn. No. 

482 : 1-366, text figs, and pis. 
Kerr, R. 1824. A general history and collection of voyages and travels. London and Edin- 
burgh, pp. vii + 503. 
Kuhn, O. 1930. Die Robben der Adria. Zool. Gart., Lpz. 2 : 194-197. 
Lafuente, Domenech. A. 1946. Algo sobre Rio de Oro. Madrid (not seen). 
Lesson, R. P. 1828. Dictionnaire Classique d'Histoire Naturelle. Paris, 13 : 1-648. 
Lewis, C. B. 1948. The West Indian seal. Nat. Hist. Notes, Jamaica, No. 34 : 169-171. 
Lobstein, J. F. 1817. Observations d'Anatomie compar6e sur le Phoque a ventre blanc. 

/. Med. Chirurgie, Pharmacie. Paris, 39 : 20-59. 
Lucas, F. A. 1891. Animals recently extinct or threatened with extermination as represented 

in the collections of the U.S. National Museum. Rep. U.S. Nat. Mus. 1889 : 609-649. 
Lungo, A. del. 1935. La Foca Monaca, Monachus albiventer, Bodd. nei mari di Sardegna. 

Rass. Faunist. 2 : 13-19. 
McMurtrie, H. 1834. Cuvier's Animal Kingdom. London, pp. xx -\- 508, illust. 
Markowski, S. 1952. The Cestodes of Pinnipeds in the Arctic and other regions. Journ. 

Helminth. 26 : 17 1-2 14. 
Matschie, P. 1905. Eine Robbe von Laysan. S. B. Ges. Naturf. Fr. Berl. 1905 : 254-262. 
Menis. 1848. II Mare Adriatico, 153. Zara, 1848 (not seen). 
Mohr, E. 1942. Tragzeitverhaltnisse der Robben. Zool. Anz. 139 : 176-183. 
Monod, Th. 1923. Note sur la presence du Monachus albiventer Bodd. sur la c6te Saharienne. 

Bull. Mus. Nat. Hist. Nat. Paris, 29 : 555-557. 

1932. Phoques Sahariens. Terre et la Vie, 12 : 257-261. 

1945. Un ordre nouveau de mammiferes pour la faune d'A.O.F. Notes Afr. 25 : 14-15, 

figs. 9-10. 

1948. Le phoque-moine dans l'Atlantique. Publ. Inst. Zool. Porto, nr. 34, 1-19, 1 pi. 

Moore, J. C. 1953. Distribution of Marine Mammals to Florida waters. Amer. Mid. Nat. 

49 : 117-158. 
Nilsson, S.,1838. Utkast till en systematisk indelning af Phocaceerna. K. Svenska Vetensk 

Akad. Handl. 1837 : 235-240. 
Olafsen, E. 1774-75. D es • • • E' Olafsens und des ... P. Povelsens Reise durch 

Island . . . beschrieben von ... E. Olafsen. Kopenhagen and Leipzig. 2 parts 

(in 1 Vol.). 
Owen, Richard. 1866. On the Anatomy of Vertebrates. 3 vol. London : Longman Green 

& Co. 
Pennant, T. 1793. History of Quadrupeds. 3rd ed., 2 : 324 pp., 190 pis. London. Seals, 

p. 266-291. 
Peron, F. 1816. Voyage de decouvertes aux Terres Australes. Paris, 2 : pp. xxxi -f 471. 
Postel, E. 1950. Un phoque tropical : le phoque moine. Nature, Paris, No. 3187 : 341-342. 
Ranzani, C. 1823. Osservazioni sulla dentatura della Foca a ventre bianco, Phoca albiventer 

Bodd. Opuscoli scientifici, 4 : 58-60. 
Reiser, O. 191 2. Uber die Erbeutung einer Monchsrobbe (Pelagius monachus) bei Fort Opus. 

Wiss. Mitt. Bosn. Herz. 12 : 665-666. 
Rondoletius, G. 1554. Libri de Piscibus Marinis. 2 vols, (in 1), illust. Lugduni. 
Sarmento, A. A. 1948. Vertebrados da Madeira. 1. Mammiferos, Aves, Repteis, Batra- 

quios. Funchal. 317 pp. 
Schauinsland, H. 1899. Drei Monate auf einer Koralleninsel {Laysan). Bremen. 
Shaw, G. 1800. General Zoology. I : pt. 2. London, pp. vii + 249-552. pis. 70-121. 
Sloane, H. 1707. A voyage to ... Jamaica. London, 2 vols., illust. 


Thompson, D'Arcy W. 1910. The Works of Aristotle. Translated into English. Oxford, 

pp. xv + 633. 
Townsend, C. H. 1906. Capture of the West Indian Seal (Monachus tropicalis) at Key West, 

Florida. Science, n.s. 23 : 583. 

1909. The West Indian seal at the aquarium. Science, n.s., 30 : 212. 

1923. The West Indian seal. /. Mammal. 4 : 55. 

Troitzky, A. 1953. Contribution a l'etude des Pinnipedes a propos de deux phoques de la 
Mediterranee ramenes de croisiere par S.A.S. le Prince Rainier III de Monaco. Bull. 
Inst. Oceanogr. Monaco, No. 1032 : 46 pp. 

Trouessart, E. L. 1910. Faune des Mammiferes d'Europe. Conspectus Mammalium Europae. 
Berlin, pp. xvii -f 266. 

True, F. W., and Lucas, F. A. 1885. On the West Indian seal {Monachus tropicalis, Gray) 
Rep. U.S. Nat. Mus. 1884 : 331-335, pis. i-iii. 

Ugolini, R. 1902. II Monachus albiventer Bodd. del Pliocene di Orciano. Palaeontographia 
Italica, 8 : 1-20, pi. iii. 

Walker, W. F. 1886. The Azores : or Western Islands. A Political, Commercial and Geo- 
graphical Account. London, pp. viii + 335- 

Ward, H. A. 1887a. The West Indian seal [Monachus tropicalis). Nature, 35 : 392. 

18876. Notes on the life history of Monachus tropicalis, the West Indian seal. Amer. 

Nat. 21 : 257-264, pi. xii. 


Monachus monachus. Photographs of the specimen from Oran in the Jardin des Plantes, 
Paris. The animal was blind in its right eye. Phot. J. E. K. 

11 JAN 1956 

Bull. B.M. [N.H.) Zool. 3, 5 




Monachus monachus. a. From Hermann, 1779. b. From Buffon, 1782. c. From 
Pennant, 1793. 

Bull. B.M. (N.H.) Zobl. 3, 5 



Monachus tropicalis. Lateral view of skeleton 1887 . 8 . 5 . 1 

hull. B.M. {N.H.) Zool. 3, 5 



M. schauinslandi. Skull of type specimen No. 32795, Zoological Museum, Berlin. A. 
Dorsal view. b. Ventral view. c. Lateral view. d. Dorsal view of lower jaw. e. Lateral 
view of lower jaw. 

hull B.M. (N.H.) Zoot. 3, 5 





M. tropical is. a. Fore limb. 

Hind limb. 

Bull. B.M. (N.H.) Zoo!. 3, 5 



Innominate bones of a. L. weddelli, b. M. monachus, c. H. grypus. 

Bull. B.M. (N.H.) Zool. 3, 5 



B C 




11 JAN 1956 



2 2 FEB 1956 


W. J. REES and G. E. MAUL 



ZOOLOGY Vol. 3 No. 6 

LONDON: 1956 


W. J. REES and G. E. MAUL 




Pp. 257 — 293; Pis. 9-10 : 4 Text-Figures 



ZOOLOGY Vol. 3 No. 6 

LONDON: 1956 

(NATURAL HISTORY), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical Series. 

Parts will appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

This paper is Vol. 3, No. 6 of the Zoological series. 


Issued February, 1956 Price Sixteen Shillings 



By W. J. REES and G. E. MAUL 


I. Introduction ..... 
II. A Summary of Previous Records 

III. Additions to the Fauna . 

IV. An Analysis of the Cephalopod Fauna 

i, The coastal fauna 

2. The benthic slope fauna 

3. The bathypelagic slope fauna 

4. The pelagic surface fauna . 

5. The epipelagic fauna 

6. The oceanic bathypelagic fauna . 

7. The abyssal fauna 

V. Cephalopoda as Food of Alepisauvus ferox 
VI. References ..... 




The cephalopod fauna of Madeira is revised and no fewer than 22 additional species are 
added to the list, chiefly on account of the examination of the stomach contents of A lepisaurus 
ferox Lowe, Aphanopus carbo Lowe and the sperm whale. A total of 18 species (many of them 
rare and seldom taken in the nets of deep sea expeditions) are recorded from Alepisauvus ferox. 
The following little known species have been found in the stomachs of sperm whales at Madeira : 
Tetronychoteuthis dussumieri (Orbigny), Lepidoteuthis grimaldi Joubin and Architeuthis sp. 
The composite fauna is analysed and compared with that of the Mediterranean. 


This preliminary paper on the cephalopod fauna of Madeira has its origins in an 
investigation of the stomach contents of the voracious fish, Alepisauvus ferox Lowe. 
The work at Madeira was pursued for some years prior to the despatch of all the 
cephalopoda taken (together with representative local collections) to the British 
Museum (Natural History) for identification. The material yielded many species 
scarce in collections and it was deemed an opportune time to produce a new list 
of Madeiran cephalopods. 

This list is rich in the names of seldom-recorded species, chiefly because the 
fishes Alepisauvus fevox, Aphanopus cavbo Lowe and the sperm whale have been 
examined for stomach contents. The occurrence of the various rare species are 
noted in the main part of the text, but special attention is drawn to the records of 
Lepidoteuthis gvimaldi Joubin, Tetvonychoteuthis dussumievi (Orbigny), Histioteuthis 
bonelliana (Ferussac) and Avchiteuthis from the sperm whale. 

zool. 3, 6. 



In this preliminary paper we have not given full descriptive accounts of the species 
found ; this is reserved for further accounts of interesting species either jointly or 


The first record of a cephalopod to be recorded in scientific literature from the 
vicinity of Madeira was of a Taonid which d'Orbigny referred to Loligopsis pavo 
Lesueur in the great Histoire des cephalopodes acetabuliferes (1835-1848) . It was later 
pointed out by Rochebrune (1884) and Hoyle (1884) that d'Orbigny had confused 
two species under this name and the Madeiran specimen (taken in sight of Madeira 
by the naturalist Dussumier) was redescribed by the former under the name Phas- 
matopsis cymoctypus. 

Robert MacAndrew, who dredged extensively on the coast of Spain and Portugal, 
both in the Atlantic and in the Mediterranean, does not mention a single cephalopod 
from his dredgings at Madeira (1849-50). 

White and Johnson, in their Handbook of Madeira (i860) list the following six 
species : 

Loligo vulgaris Lamarck. 

Sepia officinalis L. 

Octopus vulgaris Lamarck. 

Octopus cuvieri (=0. macropus Risso). 

Argonaut a argo L. 

Ocythoe tuber culata Rafinesque. 

In their second edition (1885) the fourth and last species are not listed. Hoyle 
(1885, p. 69) confirmed the occurrence of Argonauta argo at Madeira in his Challenger 

More species were added to the list by Girard (1892) who reported on the collections 
of the Abbe Ernest Schmitz of the Seminario do Funchal, adding or confirming the 
existence of the following at Madeira : 

Loligo forbesi Steenstrup. 

Spirula peronii Lamarck 1 (=Spirula spirula L.). 

Sepia filliouxii Lafont (— S. officinalis). 

Todarodes sagittatus (Lamarck). 

Ocythoe tuber culata Rafinesque. 

Octopus tuberculatus Blainville. 2 

Girard was aware of the differences between L. vulgaris and L. forbesi and as the 
species submitted to him by Schmitz proved to be L. forbesi, the former species was 
dropped from the list ; nor was he able to confirm the existence of 0. macropus in 
the area. 

1 Apparently empty shells. 

* Presumed here to be a juvenile specimen of 0. vulgaris. 


Watson (1897) added no new species to the list of Madeiran cephalopods but 
confirms that 0. vulgaris is a common species and states that Sepia officinalis is rare. 

From 1890 onwards the various deep sea expeditions have taken cephalopods in 
the N. Atlantic, sometimes near, sometimes some distance off Madeira, and the list 
compiled by Nobre (1937) includes 20 species of which several have been taken a 
considerable distance away and could not be termed local fauna. Some of the 
species which he records were taken as far west from Madeira as the African coast 
is to the east of it. The following species were taken too far away to be retained as 
local Madeiran records : 

Sepietta oweniana (Orbigny). 
Onychia caribaea Lesueur. 
Brachioteuthis riisei (Steenstrup) . 
Histioteuthis bonelliana (Ferussac). 
Taonius pavo (Lesueur) . 
Japetella diaphana Hoyle. 

As Girard's Octopus tuberculatus probably falls into the synonymy of 0. vulgaris, 
Nobre's list becomes amended to 13 cephalopods known from the immediate vicinity 
of Madeira. To these may be added Joubin's record of Leachia cyclura (1920). 


The Madeiran archipelago has a much richer cephalopod fauna as our investiga- 
tions have shown, and we do not think we have by any means exhausted the numbers 
of species which should occur off Madeira. A great many other cephalopods known 
from the central North Atlantic remain to be reported from this area. Below is 
given a list of 22 additional species which we are now recording from Madeira. 

Heteroteuthis dispar (Ruppell). 
Loligo vulgaris Lamarck. 
Abraliopsis morisii (Verany). 
Enoploteuthis leptura (Leach). 
Pyroteuthis margaritifera (Ruppell). 
Cucioteuthis unguiculata (Molina). 
Tetronychoteuthis dussumieri (Orbigny). 1 
Lepidoteuthis grimaldi Joubin. 1 
Onychoteuthis banksi (Leach). 
Architeuthis sp. 

Histioteuthis bonelliana (Ferussac). 
Ommastrephes pteropus Steenstrup. 
Thysanoteuthis rhombus Troschel. 
Chiroteuthis veranyi (Ferussac). 
Mastigoteuthis schmidti Degner. 

1 These two species are here recorded from the stomachs of sperm whales taken at Madeira. It is 
hoped to provide a full description in another paper, 


Taonius pavo (Lesueur) . 
Cranchia scabra Leach. 
Japetella diaphana Hoyle. 
Eledonella pygmaea Verrill. 
Vitreledonella richardi Joubin. 
Alloposus mollis Verrill. 
Tremoctopus violaceus delle Chiaje. 

Family Spirulidae 
Spirula spirula L. 

Spirula peronii, Girard, 1892, p. 219 ; Watson, 1897, p. 318, Nobre, 1937, P- 5- 

This species is known only from the dead shells at Madeira ; these are cast up 
on the beaches of Porto Santo occasionally and are also known from Canigal (Girard) . 
Bruun (1943) reporting on the living Spirula captured by the " Dana " expeditions, 
believed that this species inhabits the continental slope, living at depths not 
exceeding 1750 metres. 

Family Sepiidae 
Sepia officinalis L. 

Sepia officinalis, White & Johnson, i860 ; Johnson, 1885 ; Watson, 1897, P- 3 J 6 ; Nobre, 1937, 

P. 5- 

Sepia fillouxii, Girard, 1892, p. 219. 

Material seen : 

1 specimen, dorsal mantle length of 205 mm. (Maul collection No. 144). 

According to Watson (1897), Sepia officinalis is rare at Madeira, but the species is 
very common all the year round at Madeira and adult specimens measuring about 
25 cm. (in dorsal mantle length) are often seen at the local fish market. The species 
goes by the name of Chouco and is eaten by the local population. 

Family Sepiolidae 

Heteroteuthis dispar (Riippell) 

Material seen : All except No. 3281 from the stomachs of Alepisaurus ferox. 

No. 3281, 1 mutilated specimen, dorsal mantle length 20 mm. from the 

stomach of Aphanopus carbo, io.i.51. 
No. 3365, 1 specimen, dorsal mantle length 25 mm., 10. iv. 1952. 
No. 3490, 4 specimens, dorsal mantle lengths of 25, 28, 29 and 20 mm. — no 

No. 3802, 1 specimen, dorsal mantle length of 22 mm. — no date. 
No. 3848, 2 specimens, dorsal mantle lengths of 8 and 12 mm. — no date. 

This sepiolid has not previously been taken at Madeira, and although reported 
from numerous stations in the Mediterranean (Degner, 1925). it has been found on 
comparatively few occasions in the Atlantic. Each major expedition has, however, 


secured specimens ; those found nearest to Madeira being those captured by the 
" Talisman " (29 2' N., 14 49' N.) between the Canaries and the African coast 
(Fischer & Joubin, 1907), and those of the " Thor " from just outside the Straits of 
Gibraltar (Degner, 1925). Heteroteuthis dispar is often regarded as a deep sea 
cephalopod, but it is difficult to justify this claim, which seems to be based on the 
fact that the species has a luminous secretion. To quote Harvey (1952, p. 283) : 
"It is sometimes said that the luminous secretion of deep sea Heteroteuthis has 
replaced the ink of surface dwellers. This black fluid forms a poisonous smoke 
screen which confounds the enemy while the squid makes a getaway. Although it 
is not true that the luminous secretion has completely replaced the ink, it may 
serve a similar purpose — to draw the attention of the enemy to a mass of lighting 
material, while the squid moves off in another direction." However, an examination 
of Degner' s details of capture in the Mediterranean and Atlantic shows that captures 
were more frequent with only 300 metres of wire out than in deeper hauls, and it can 
be reasonably assumed that the net was fishing at only half or less than half this 
depth. The fact that Heteroteuthis has retained its ink sac is also an indication 
that it can and does live above the threshold of light. Whether it can also live 
below the threshold of light is so far unknown. As an item in the food of Alepi- 
saurus ferox, it probably occurs at the depths at which this fish is taken on tunny 
hooks at Madeira. It is also a regular item in the diet of the long-finned albacore 
(Germo alalunga) in the Bay of Biscay (Bouxin & Legendre, 1936, p. 36), and it has 
been reported from the stomach of the dolphin (Delphinus delphis) by Joubin 
(1900, p. 10). 

Sepietta oweniana (Orbigny) 

Sepiola scandica, Fischer & Joubin, 1906, p. 204. 

Sepietta oweniana, Bouxin & Legendre, 1936, p. 32 ; Nobre, 1937, P- 5- 

Material seen : 

No. 2812, 2 specimens, dorsal mantle lengths of 12 and 17 mm., Funchal 
Harbour, 12. hi. 1943. 

The " Travailleur " took this species in position 32 40' N., 18 54' W., some 
distance to the south-west of Madeira. Its occurrence at Madeira is confirmed from 
two specimens we have seen. 

Family Loliginidae 

Loligo vulgaris Lamarck 
Material seen : 

No. 2966, 1 juvenile, no data. 

No. 3164, 5 juveniles from Funchal Harbour, 4. v. 1950. 
No. 3848, 1 juvenile, dorsal mantle length of 7 mm. from stomach of Alepi- 
saurus ferox. 

The specimens from Funchal Harbour are juveniles of only 30-35 mm. in dorsal 
mantle length. Loligo vulgaris is recorded as far north as the coast of Denmark, 


but it is doubtful whether it breeds anywhere north of the latitude of Ushant at 
the mouth of the English Channel. As a Lusitanian species it is well known in the 
Mediterranean and on the Mauritanian coast, while recently Adam (1952) has demon- 
strated that it occurs off the Congo and as far south as Algoa Bay. It has been 
reported from the Canary Islands (Odhner, 1931) and, although said to occur at the 
Azores (Drouet, 1858), its presence there requires confirmation. 

Loligo forbesi Steenstrup 
Loligo forbesii, Girard, 1892, p. 219 ; Nobre, 1937, P- 6. 

No. 4355, Funchal Market 1 $ of 270 mm. in dorsal mantle length. 

Girard based his identification of this species on many examples sent to him by 
the Abbe Ernest Schmitz of Funchal. This species has often been reported from 
the neighbouring African coast. Locally, Loligo goes by the name of Lula da costa, 
but it is not known whether the two species are confused under this name. It is 
very common in winter, and as an article of food stands next to 0. pteropus in 
importance. Market specimens range from about 200 mm. to 500 mm. in dorsal 
mantle length. 

Abralia veranyi (Riippell) 

Abralia veranyi, Berry, 1926, p. 257. 
Abralia veranyi, Nobre, 1937, P- 7- 

Material seen : 

No. 2664, 1 specimen of 47 mm. dorsal mantle length, Funchal Harbour, 

at night, 16.X.1941. 
No. 2964, no data — 5 specimens, 37-50 mm. in dorsal mantle length. 
No. 3514, Funchal Museum (no date). 

This luminous squid is a Mediterranean-Atlantic species ; most of the records 
of its occurrence are from the Mediterranean (Messina, Nice, Genoa, Naples and 
Toulon), while in the Atlantic (apart from Madeira) it has been found at compara- 
tively few places, viz : Cockburn Town, San Salvador Island (Adam, 1941) and off 
the Congo (Adam, 1952). 

These specimens provide additional records for Madeira. No. 3514, a male, has 
the left ventral arm hectocotylized just as figured by Adam (1952, p. 63, fig. 25E). 
This specimen has a dorsal mantle length of 42 mm. One of the female specimens 
(Lot No. 2964) has a dorsal mantle length of 50 mm. ; this is considerably bigger 
than Adam's largest specimens, which were up to about 40 mm. in dorsal length 
(Adam, 1952, p. 70). 

Abraliopsis morisii (Verany) 
Material seen : 

No. 3848, 6 specimens of 8-25 mm. in dorsal mantle length from the stomach 
of Alepisaurus ferox. 


This is the first time this small luminous squid has been recorded from Madeira 
and from the stomach of Alepisaurus ferox. Records of this species from the 
Mediterranean and the Atlantic are fairly numerous (see Chun, 1910). 

Enoploteuthis leptura (Leach) 
Material seen : 

No. 2664, Funchal Harbour at night, 16.X.1941, 1 specimen. 

No. 2963, Madeira, 1 specimen, no data. 

No. 3487, Funchal Harbour at night,, 1 specimen. 

These are the first Madeiran records of this rather scarce cephalopod. It is known 
from the African coast and Pacific (Ferussac & d'Orbigny, 1 835-1 848) and a juvenile 
from the Atlantic (position o° 29' N., 18 57' W.) was captured by the Deutsche 
Sudpolar-Expedition and figured by Chun (1910, Taf. xl, figs. 5 and 6) in the 
" Valdivia " report. 

Pyroteuthis margaritifera (Ruppell) 
Material seen : 

No. 2961, 2 specimens, dorsal mantle length of 35 mm., from stomach of 
Alepisaurus ferox, March and April, 1944. 

This is another well known Mediterranean- Atlantic, luminous cephalopod. 
Joubin (1924, p. 52) reported it from the Azores but it does not seem to have been 
recorded from Madeira, neither has it been recorded as an item in the diet of Alepi- 
saurus ferox. The " Thor " took specimens in the Eastern Mediterranean with 
200-300 metres of wire out, so that the species can be assumed to exist up to 100 
metres of the surface. 

Family Octopodoteuthidae 
Cucioteuthis unguiculata (Molina) 
Cucioteuthis unguiculata, Joubin, 1900, p. 51 ; Nobre, 1937, P- 6. 

The " Princess Alice " took this species in position 32 32' 10" N., 19 24' 40" W., 
to the west of Madeira. There is a large brachial crown, in the Municipal Museum, 
Funchal, from a specimen which was found dead floating at the surface near the 
shore to the west of Funchal. 

Onychia caribaea Lesueur 

Teleoteuthis jattai Joubin, 1900, p. 67. 
Teleoteuthis caribaea, Nobre, 1937, P- 7- 

Nobre includes this species in his list of Madeiran cephalopods, but the position 
at which this species was taken by the " Princess Alice " (1897) Cruise, St. 812, 
31 04' N., 27 11/ W.) is nearly as far to the south-west of Madeira as these islands 
are from the African coast. The species must therefore be omitted from the list. 


Onychoteuthis banski (Leach) 
Material seen : 

No. 2455, no data, 1 specimen of 100 mm. in dorsal mantle length. 
No. 3507, no data, 1 specimen of 130 mm. in dorsal mantle length. 
No. 2965, no data, 1 poorly preserved specimen of 35 mm. dorsal mantle 

No. 3571, 1 specimen of 48 mm. dorsal mantle length, from stomach of 

Alepisaurus ferox, 4 . xii . 1952. 
No. 3848, 45 specimens, dorsal mantle lengths of 10, 13, 21-24 mm., stomach 

of Alepisaurus ferox. 
No. 3509, 1 mutilated juvenile, dorsal mantle length 24 mm., from the 

stomach of Alepisaurus ferox. 

Onychoteuthis banksi is here recorded for the first time from Madeira. No. 2965 
is in poor condition and only provisionally referred to this species. 

This is a well-known oceanic surface species which, in the neighbourhood of 
Madeira, was taken in position 30 54' N., 24 11' N., by the Prince of Monaco in his 
1912 cruise (Joubin, 1924, p. 48). Other nearby records for the " Terra Nova " 
and the " Ara " are given by Rees (1949, p. 43) who drew attention to the habit of 
" flying " in this species. New records of specimens being found on the deck of 
ships are given by Adam (1952, p. 77). 

Family Architeuthidae 

Architeuthis sp. 

A sperm whale, harpooned off Sao Lourenco on 12th June, 1952, vomited a squid 
which had evidently just been swallowed because the latter still exhibited some signs 
of life. This squid was large, with a weight of 150 kilograms, and could only have 
been a species of Architeuthis. The following measurements were taken : 

Overall length (i.e. including tentacles) . . 10,600 mm. 

Dorsal mantle length . . . . .1,860 

Length of tentacles 
Length of arm 
Circumference of mantle 
Circumference of tentacles 





Architeuthis was taken by the French steamer " Alecton " between Madeira and 
Teneriffe in November, i860, and the species was subsequently given the name 
Loligo bouyeri by Crosse and Fischer (1862, p. 138). A number of species of Archi- 
teuthis have been described since Verrill's classical monograph and it is by no means 
certain how many kinds there are. Fragments of the above specimen are kept in 
the Museu Municipal, Funchal, and it is hoped to present a more detailed description 
in another paper. 


Family Histioteuthidae 
Histioteuthis bonelliana (Ferussac) 

Histioteuthis ruppelli, Joubin, 1900, p. 98. 
Histioteuthis bonelliana, Nobre, 1937, P- 6. 

Material seen : 

No. 3905, 1 specimen, dorsal mantle length of 53 mm., from stomach of 
Aphanopus carbo, 30 . x . 1953. 

Nobre records this species as Madeiran on the basis of an arm found in position 
41 o' N., 12 15' W., by the " Princess Alice " in 1894 (Joubin) ; this position is off 
the north coast of Portugal, so that this could not be justified as a Madeiran record 
of this species. Histioteuthis does occur, however, around Madeira, as recorded 
above, and we are indebted to Mr. H. M. Sieyes for a photograph of a large brachial 
crown of this species found in the stomach of a sperm whale captured off Sao Laurenco 
lighthouse, Madeira on 12th June, 1952. It is interesting to note that this sperm 
whale feeds on this species at Madeira, as at the Azores and elsewhere. The speci- 
men from the stomach of Aphanopus carbo was much smaller, with an overall length 
length of about 170 mm. It is noteworthy that Verrill (1880, p. 234) recorded 
H. bonelliana from Alepisaurus ferox (as H. collinsii). 

Family Brachioteuthidae 
Brachioteuthis riisei (Steenstrup) 

Tracheloteuthis riisei, Joubin, 1924, p. 75. 
Brachioteuthis {Tracheloteuthis) riisei, Nobre, 1937, P- 6. 

This wide-ranging surface cephalopod is included in the Madeiran fauna by Nobre 
on the strength of a record of capture of this species in position 31 45' N., 20 17' W., 
by the " Princess Alice " (Joubin). This is some distance south and west of Madeira, 
but such a widely distributed species is almost certain to be found in this area. 

Family Ommastrephidae 
Todarodes sagittatus (Lamarck) 

Todarodes sagittatus, Girard, 1892, p. 220. 
Ommastrephes sagittatus, Nobre, 1937, P- 8. 

Material seen : 

No. 3355, 1 juvenile, dorsal mantle length of 83 mm. from the stomach of 

Alepisaurus ferox, 8 . iv . 1952. 
No. 3356, 1 juvenile, dorsal mantle length 86 mm. from the stomach of 

Alepisaurus ferox, 8 . iv . 1952. 
No. 3363, 1 juvenile, dorsal mantle length 85 mm. from the stomach of 

Alepisaurus ferox. 
No. 4347, Funchal Market, i specimen of 185 mm. in dorsal mantle length. 


Madeiran records in the literature are few and are based on the following speci- 
mens : i $ taken by Schmitz, i $ by Nunes. To these may be added the specimens 
from the stomach of Alepisaurus ferox noted above. 

The sucker rings of the tentacles are mostly absent : the very long-sucker-bearing 
region of the tentacle, extending for the greater part of its length, the almost smooth, 
large tentacular rings and the presence of four rows of suckers near the tip of the 
tentacles indicate that the species is T. sagittatus. Locally Todarodes is called 
Cartucho ; it is extremely common on the fishing grounds in March, April and May. 
It is also taken during other months, but it is rare at these times. 

As a common cephalopod on the fishing grounds of the eastern North Atlantic 
and the Mediterranean, Todarodes is preyed on by a number of animals, including 
the dolphin, Delphinus delphis (Joubin, 1895, p. 32), Germo alahinga (Bouxin and 
Legendre, 1936) and Gadus calliarias {Michael Sars Stations 7, 86, 352 and 115). 
Joubin (1895, 1900, 1920 and 1924) also reported various ommastrephid fragments, 
some of which probably belong to this species, from Polyprion americanum and 
Grampus griseus. 

Ommastrephes pteropus Steenstrup 
Material seen : 

Nos. 65 and 66, 2 fragments, Funchal Harbour, 16.iv.1941. 

No. 193, 1 juvenile, dorsal mantle length 58 mm., 10. v. 1940. 

No. 3443, 1 specimen, dorsal mantle length 270 mm. — no data. 

No. 3678, 1 juvenile, dorsal mantle length 50 mm. from stomach of Alepi- 
saurus ferox, 25 . iii . 1953. 

D. W. Tucker Collection, No. 605, 1 juvenile, dorsal mantle length 80 mm. 

D. W. Tucker Collection, Nos. 639-644, 6 half-grown specimens, gaffed 
at night on Madeiran fishing grounds, August, 1953. 

All these specimens have been referred to 0. pteropus ; some of them might equally 
well have been referred to 0. bartrami as we find that the diagnostic key given by 
Pfeffer (1912, p. 465) is unreliable and our specimens clearly belong to one species. 

Adam (1952, p. no) also noted the unreliable nature of the connective apparatus 
on the tentacles as a means of separating bartrami from pteropus. The number of 
suckers, proximal to the nearest connective button, will vary as new buttons develop 
with the growth of the tentacle, as indicated in one of our larger specimens where 
there is a rudimentary button ; this points to the great need for a revision of these 
species with more material than we have at our disposal. 

The relationships of these two species to Ommastrephes caroli is equally puzzling. 
In the species caroli the lateral membrane is a broad triangular flap in adult speci- 
mens (Rees, 1950, pi. 1) while in younger specimens it is much narrower (see Hertling, 
1938). Our specimens from Madeira (those of 2-3 ft. in total length) possess narrow 
lateral membranes on the third arms which might conceivably develop into the 
characteristic paper-thin flaps of caroli. As in the latter their chromatophores are 
closely set and a deep chestnut-brown in colour, 


Ommastrephes pteropus is very abundant during the summer months of July, 
August and September and is extensively used for baiting the tackle for Aphanopus 
carbo, tunny and other fish. It is found in great numbers in locally caught Germo 
obesus. Large quantities of this squid, which goes under the local name of 
Pota de limao or Lula de limao, are sold in the fish market for consumption by 
the local people, who greatly appreciate it as food. The large, lemon-coloured 
dorsal mark has been observed to flare up with a bright light when the animal 
is gaffed. 

Ommastrephids : 

No. 2957, 1 juvenile, dorsal mantle length of 50 mm. from stomach of Alepi- 

saurus ferox, March-April, 1944. 
No. 3848, 3 juveniles, dorsal mantle length of 17, 20 and 25 mm. from the 

stomach of Alepisaur us ferox. 
No. 193, 1 juvenile, dorsal mantle length of 65 mm. from the stomach of 

Alepisaurus ferox, 10 . v . 1940. 

These fragmentary specimens are in poor condition and we have not been able to 
identify them. 

Thysanoteuthis rhombus Troschel 
Material seen : 

1 specimen, dorsal mantle length of 350 mm., taken at Funchal. 

This is the first record of this powerful surface species from Madeira. Some years 
ago a school of about 20 came close inshore, and this is the only specimen it was 
possible to save for examination. Little is known about its distribution, but it 
occurs in the Mediterranean (Naef, 1 923-1 928) and has been reported from the Cape 
of Good Hope by Barnard (1934). It is likely, however, to be a cosmopolitan warm- 
water species, as Sasaki (1929) has recorded it from a number of localities in Japanese 

Chiroteuthis veranyi (Ferussac) 
Material seen : 

No. 3685, 1 Doratopsis larva of 80 mm. in overall length. Coll. A. A. Nunes, 

This larva was taken at the Pontinha at the surface at night in the light of an 
electric lamp suspended over the water. 

It is a well-known Mediterranean- Atlantic species whose larva is frequently taken 
at places like Messina and Villefranche, where there is upwelling from deep water. 
The closely related species Doratopsis exophthalmica was taken by the " Valdivia" 
in position 31 59' N., 15 5' W. to the south-east of Madeira. 

Published records of C. veranyi from the North Atlantic are few (see Adam, 1952, 
p. 111), but to judge from the number of specimens recovered from the stomachs 
of germon fished in the Bay of Biscay the species is not as rare as the absence of 
records suggests (Bouxin and Legendre, 1936). 



Mastigoteuthis schmidti Degner 
Material seen : 

1 specimen, dorsal mantle length of 107 mm., from the stomach of Alepi- 
saurus ferox, December, 1944. 

There are no previous records of Mastigoteuthis from Madeira and this specimen 
is only provisionally referred to M. schmidti because we recognize the need for a 
complete revision of the large number of species described from the Atlantic. As 
regards the proportions of mantle and fin the species is very similar to M. schmidti. 

Family Cranchiidae 

Leachia cyclura Lesueur 

Leachia cyclura, Joubin, 1920, p. 68. 

Material seen : 

No. 3513, 1 specimen, overall length 145 mm. from stomach of Alepisaurus 

ferox — no date. 
No. 3848, 1 juvenile, dorsal mantle length of 23 mm. from stomach of Alepi- 
saurus ferox. 

Leachia cyclura was taken by the Prince of Monaco on the fishing grounds of 
Madeira in 1901 (Stn. 1235, 8th September, 32 34' N., 17 45' W.). This species 
has rarely been captured in the Atlantic, and since it was described from the Atlantic 
coast of N. America by Lesueur (1821), it has been recorded by Joubin only from 
Madeira, the Azores and the Sargasso Sea. It is known from other oceans also. 

Cranchia scabra Leach 

No. 34970, 1 specimen, overall length of no mm. from stomach of Alepisaurus 

No. 34976, 1 specimen, Funchal Harbour, at night. 
No. 3497c, 1 specimen, overall length 105 mm. from stomach of Alepisaurus 


This species is well known from the sub-tropical and tropical areas of the Atlantic 
and other oceans, but does not appear to have been reported from Madeira. It is 
also recorded for the first time from the stomach of Alepisaurus ferox. 

There is no particular reason for supposing that this species is a deep-water 
animal. Its presence in the upper 200 metres has been proved by the Deutsche 
Sudpolar Expedition, which took specimens at 20 and 40 metres depth. It should 
be noted that records from 2,000 and 3,500 metres by the " Valdivia " were made 
with open nets (Chun, 1910). 


Taonius pavo (Lesueur) 

Taonius pavo, Joubin, 1900, p. 106 ; Nobre, 1937, P- 8. 

Material seen : 

No. 3902, 1 specimen without tentacles, partly macerated, from the stomach 
of Alepisaurus ferox, 26 . xi . 1953. 

This specimen agrees favourably with the specimen figured by Joubin (1900, p. 106, 
pi. viii and ix), but the colour is of a deep reddish purple. Our specimen has a 
dorsal mantle length of 275 mm. and a total length of 385 mm. 

This example, from the stomach of A. ferox, is the first to be found at Madeira. 
The specimen taken by the Prince of Monaco in 1897 (Stn. 817, position 30 42' N., 
27 32' W. was found well out in the Atlantic and a considerable distance from 

Taonius pavo has all the appearance of being a bathypelagic species, and we have 
wondered whether the occurrence of this squid (and also some others, notably, 
Mastigoteuthis schmidti and Eledonella pygmaea) in this fish can be explained by 
local up welling of deep water bringing these species near the surface. 

Taonius cymoctypus (Rochebrune) 

Loligop sis pavo, Orbigny, 1835-1849, p. 321 (pars). 
Phasmatopsis cymoctypus Rochebrune, 1884, p. 15, pi. 1. 
Taonius pavo, Girard, 1892, p. 220 ; Nobre, 1937, P- 8. 

The only specimen known of this cephalopod was taken within sight of Madeira 
by the French naturalist Dussumier and was confused with T. pavo by d' Orbigny. 
Later Rochebrune demonstrated that it was a distinct species. A new description 
of this curious squid is greatly needed. 


Family Bolitaenidae 

Japetella diaphana Hoyle 

Japetella diaphana, Thore, 1949, p. 23, fig. 14. 
Bolitaenella diaphana, Nobre, 1937, P- 3- 

Material seen : 

No. 2958, 4 specimens, total length ca. 50 mm., stomach of Alepisaurus ferox, 

March- April, 1944. 
No. 3357, 1 specimen, total length ca. 50 mm., from stomach of Alepisaurus 

ferox, 8.1V.1952. 
No. 3495, 2 specimens, total length of 50 and 70 mm., stomach of Alepisaurus 

No. 3496, 1 specimen, total length 70 mm., ? stomach of Alepisaurus ferox. 


No. 3569, 2 specimens, total lengths 55 and 80 mm., stomach of Alepisaurus 

ferox, 4.XH.1952. 
No. 3594, 1 specimen, total length 65-70 mm., stomach of Alepisaurus ferox, 

No. 3903, 1 damaged specimen, total length 85 mm., stomach of Alepisaurus 

ferox, 26.x. 1953. 

There are numerous records of this pelagic octopod in the North Atlantic, in the 
triangle, Canaries, Gibraltar, Azores (Thore, 1949, fig. 14, p. 23), but the first positive 
records from Madeiran fishing grounds are given above. 

Bouxin & Legendre (1936, p. 12) reported finding this species (one specimen in 
each of three stomachs) in the germon (Germo alalunga) captured in the Bay of 
Biscay. It is now recorded for the first time from Alepisaurus ferox and appears to 
be a regular item in the diet of this fish. 

For the Bolitaenella diaphana of Nobre, see Eledonella pygmaea. 

Eledonella pygmaea Verrill 

Eledonella diaphana, Joubin, 1900, p. 37, pi. 2, figs. 5-7. 

Material seen : 

No. 3800, 1 $ total length 150 mm., from the stomach of Alepisaurus ferox, no 

Eledonella pygmaea is less common than the preceding species. It has not 
previously been recorded from Madeira or from the stomach of Alepisaurus ferox. 

The " Princess Alice " (Joubin, 1900) took this species between Madeira and the 
African coast and there are more recent records of specimens taken by the " Dana " 
to the west and north of the Madeiran archipelago (Thore, 1949, fig. 41, p. 49). 

Family Vitreledonellidae 

Vitreledonella richardi Joubin 

Vitv eledonella alberti Joubin, 1924, p. 38 ; Nobre, 1937, P- 3- 

This species had been included in a list of Madeiran cephalopods by Nobre (1937), 
who quotes Joubin's record of a specimen taken to the N.W. of Madeira (position, 
33 40' N. to 33 52' N., 19 W. to 19 16' W.) by the Prince of Monaco. 

Family Octopodidae 
Octopus vulgaris Lamarck 

Octopus vulgaris, White and Johnson, i860 ; Johnson, 1885 ; Girard, 1892, p. 218 ; Watson, 

1897, P- 2 9° «' Nobre, 1937, P- 3- 
Octopus tuberculatus , Girard, 1892, p. 218 ; Nobre, 1937, P- 4- 
Octopus rugosus, Robson, 1929. 



Material seen : 
British Museum : 

B.M. No. 1898. 5. 10. 2, Porto Santo, Madeira, Coll. H. S. Wellcombe (Robson, 

1929, as 0. rugosus). 
B.M. No. 1912.12.31.118-119, Porto Santo, Madeira, 60 fms., R. Kirk- 

patrick (Robson, 1929, as 0. rugosus). 
Funchal Market from mouth of Conger conger, 18 . viii . 1953, Coll. D. W. Tucker. 
Funchal from a tide pool, 28. viii. 1953, Coll. D. W. Tucker. 
Funchal Museum : 

No. 2460, 1 post-larva. 

No. 140, 1 specimen, dorsal mantle length 90 mm. 

No. 3287, from the stomach of Aphanopus carbo.* 

The common octopus, which goes by the local name of polvo, is very common all 
the year round at Madeira and is on sale in the fish market. The fact that it is eaten 
by the local population may account for the scarcity of specimens available for 
examination. It is found, too, at the Cape Verde Islands, the Canaries and the 
Azores. Juvenile octopods mentioned under the name Octopus tuberculatus seem 
to belong to 0. vulgaris. 

Octopus macropus Risso 
Octopus cuvieri, Girard, 1892, p. 219 ; White & Johnson, i860 ; Nobre, 1937, P- 4- 
Material seen : 

Funchal Museum, — 

No. 39, 1 $, total length 475 mm., Camara de Lobos, 30. xi. 1940. 

No. 139, 1 <J, total length 1,010 mm. — no date. 

No. 2460, 2 juveniles in alder i stage — no date. 

No. 3390, 1 J, total length 550 mm. — no date. 

No. 3493, 1 juvenile in the alderi stage, from the stomach of Alepisaurus ferox. 

This species was originally listed as occurring at Madeira in White & Johnson's 
Handbook, but the name was omitted in the second edition (1885). Subsequent 
authors have not recorded new material, so that evidence of its occurrence at 
Madeira was needed. The new records given above dispel this doubt and confirm 
its presence at Madeira. 

Octopus sp. 

Material seen : 

No. 2960, 1 partly-digested specimen with a total length of 120 mm., from 

the stomach of Alepisaurus ferox, March-April, 1944. 
No. 3504, 1 specimen from stomach of Conger conger (L.), 18.vii.1952. 
No. 3577, 1 specimen with a dorsal mantle length of 14 mm., from stomach of 
Alepisaurus ferox, 10 . xii . 1952. 
It has not been possible to positively identify these specimens with either of the 
foregoing species or any other, because of their immaturity and poor condition. 

* From the stomach of Aphanopus carbo according to a fish gutter 
zool. 3, 6. 20 


Family Alloposidae 
Alloposus mollis Verrill 
Material seen : 

No. 2460, 1 ?, total length 115 mm., March- April, 1944. 

No. 2960, 1$, total length 115 mm., stomach of Alepisaurus ferox. 

No. 3491, 5 ?, total lengths of 80, 115, 115, 115 and 117 mm., from stomach 

of Alepisaurus ferox, no date. 
No. 3494, 1 ?, total length 175 mm., stomach of Alepisaurus ferox, no date. 
No. 3804, 1 $, total length 115 mm., stomach of Alepisaurus ferox, 12 .vi. 1953. 
No. 3848, 1 $, total length of 50 mm., stomach of Alepisaurus ferox , no date. 

This species has not been previously recorded from Madeira, but has been taken 
at the Azores and other places in the North Atlantic, mainly in proximity to coastal 
waters. Thore, who plotted its distribution (1949, fig. 69, p. 72), regards it as a 
cosmopolitan, tropical and sub-tropical coast-loving species. 

The series now reported from Madeira are all small females (50-175 mm. in total 
length, compared with VerruTs specimen of 812 mm.) (Verrill, 1882). The paucity 
of records of Alloposus are noted by Thore, who suggests that either it is a rare 
animal or is able to evade plankton nets or spends " relatively short periods of its 
life-cycle in open waters, then soon returning to a life at the bottom, especially on 
the continental slopes." These new records suggest that the species is by no means 
as rare as the literature suggests and that we lack the gear to take it in numbers. 
At the smaller sizes (50-175 mm.) it seems to be a regular item in the diet of Alepi- 
saurus ferox. 

Family Tremoctopodidae 

Tremoctopus violaceus delle Chiaje 
Material seen : 

No. 3404, 1 9, total length 480 mm. 
No. 3406, 1 9, total length 510 mm. 
No. 3408, 1 9, total length 305 mm. 

There are no previous records of Tremoctopus from Madeira. It is well known 
from the Mediterranean but records from the North Atlantic are remarkably few 
(these are summarized by Robson, 1932, and Salisbury, 1953). 

Family Ocythoidae 
Ocythoe tuberculata Rafinesque 

Ocythoe tuberculata, Girard, 1892, p. 218 ; Robson, 1932, p. 202. 
Oxythoe tuberculata Nobre, 1937, P- 4- 

Material seen : 

1 9, British Museum, No. 1858. 3. 31. 6 (Robson, 1932). 

No. 2460, 1 9, total length 115 mm. (33 mm. DML), no data. 



No. 3440, i 9, dorsal mantle length of 190 mm. — no data. 
No. 3441, 1 ?, dorsal mantle length of 180 mm. — no data. 
No. 3492, 1 ?, dorsal mantle length of 50 mm., from the stomach of Alepi- 

saitrus ferox. 
No. 3848, 1 ?, dorsal mantle length of 12 mm., from the stomach of Alepi- 

saurus ferox. 

Girard records having seen a specimen of this species in the Museu do Seminario 
do Funchal and there is another in the British Museum. 

There are two large specimens noted here (Nos. 3440 and 3441) ; They compare 
favourably in size with one reported by Berry (1916) from California. Large as 
these are, Robson (1932, p. 205) records having seen one of 280 mm. (DML) at 

The young ? specimen (No. 3492) has a total length of 195 mm., and is remarkably 
well preserved. The tubercules, connected by ridges, on the ventral side of the 
mantle, are well developed and the aquiferous pores prominent. In another, even 
smaller specimen (No. 2460), these characteristics are easily seen. 

Ocythoe tuberculoid is another cosmopolitan species in tropical and sub-tropical 
waters of all oceans ; as far as we know, it is essentially a species which frequents 
the surface layers. Bouxin & Legendre (1936, p. p. 31) found it in the stomach of 
the germon in the Bay of Biscay, while Joubin (1900, p. 26) reported three females 
from the stomach of Grampus griseus taken off Monaco. 

It is now recorded for the first time as an item in the diet of Alepisaurus ferox. 

Family Argonautidae 
Argonauta argo L. 

Argonauta argo, Girard, 1892, p. 218 ; Hoyle, 1885, p. 69 ; Watson, 1897, p. 274 ; Nobre, 

1937- P- 5- 

Material seen : 

No. 3489, 1 juvenile $, dorsal mantle length 37 mm., no data. 

Early records from Madeira and Porto Santo are noted by Girard and Hoyle 
respectively, and it is not surprising that this cosmopolitan, warm-water species 
should be found here. 

A juvenile female (without shell) is provisionally referred to this species. Watson 
(1897, p. 274) records one perfect specimen from Porto Santo, and by this we assume 
(as he was a conchologist) a shell or brood chamber. 

Females of about 20-100 mm. in dorsal mantle length are found in great quantities 
in the stomachs of Alepisaurus ferox at Madeira during the months of March, April 
and May. There is one record for February and another of a single specimen in 
November. The shells are usually crushed, but sometimes they are quite intact. 
Eggs have been seen in shells of not more than 50 mm. in diameter. There is, in 
addition, a single record of a small female (of about 35 mm. in shell diameter) caught 
in Funchal Harbour at night. Large empty shells are not uncommonly found 
washed up on the beach of Porto Santo. 



Surrounded as it is by deep water, the Madeiran Archipelago has a composite 
fauna with elements from different habitats. In general, the fauna may be said 
to be a Mediterranean- Atlantic one, which we group as follows : 

1. The coastal fauna 

This includes both littoral and continental shelf species, viz : Sepia officinalis, 
Sepietta oweniana, Loligo vulgaris, Loligo forbesi , Todarodes sagittatus, Octopus vulgaris 
and 0. macropus. 

All these are common also in the Mediterranean and eastern Atlantic. Only the 
two species of Octopus have a wider distribution in the Atlantic and elsewhere. 

2. The benthic slope fauna 

No Madeiran representatives of this fauna have yet been taken. The deep-water 
octopods of the genus Bathypolypus are typical examples of species found on the 
bottom below the 200 m. line. 

3. The bathy pelagic slope fauna 

In the grouping of these species we are on less sure ground because of the scarcity 
of locality and depth records. Despite this we believe that the normal habitat of the 
adults of the species enumerated below is close to the bottom on the continental 
slope below the 200 m. line, with a lower limit perhaps at 2,000 m. 

Spirula spirula. 
Lepidoteuthis grimaldi. 
Cucioteuthis unguiculata. 
Tetronychoteuthis dussumieri . 
Architeuthis sp. 
Histioteuthis bonelliana. 
Ommastrephes pteropus. 

4. The pelagic surface fauna 

By these we mean the species that are habitually found on or very close to the 
surface. They include : 

Ommastrephes pteropus (juveniles). 
Onychoteuthis banksi. 
Tremoctopus violaceus. 
Ocythoe tuber culata. 
Argonauta argo. 
And possibly also 

Thysanoteuthis rhombus. 


5. The epipelagic fauna 

In grouping pelagic forms we have to remember that many species have epipelagic 
larvae while the adults may live well below the threshold of light. Up welling from 
deep water on some coasts and near oceanic islands sometimes brings charac- 
teristically deep-water species to the surface and it is not always possible to reach 
positive conclusions. 

The following oceanic species appear to live in the upper 500 metres : 

Heteroteuthis dispar. 
Abralia veranyi. 
? Enoploteuthis leptura. 
Pyroteuthis margaritifera. 
Chiroteuthis veranyi (? juveniles only). 
Leachia cyclura (juveniles). 
Cranchia scabra. 
Japetella diaphana (juveniles). 
Alloposus mollis (some part of its life history). 

6. The oceanic bathy pelagic fauna 

These include deep-water species which live at or below the threshold of light (ca. 
700 m.) and which are not confined to the immediate vicinity of the continental 
slope. These species often have larvae which are epipelagic. 

Chiroteuthis veranyi (adult). 
Mastigoteuthis schmidti. 
Leachia cyclura (adult). 
Taonius pavo. 
Japetella diaphana. 
Eledonella pygmaea. 
Vitreledonella richardi. 

We think that the occurrence of these species in the Madeiran list is probably due 
to the fact that there is upwelling which brings them into the upper waters or into 
the hunting grounds of Alepisaurus. 

7. The abyssal fauna 

Truly abyssal forms like V ampyroteuthis infernalis Chun have not been taken at 

In drawing up these lists we have paid but little attention to early records made 
with open nets from deep water. All too often when a plankton haul has been made 
with an open net from great depths (say 3,000 m.) to the surface, the resulting catch 
is reported as from 3,000 metres. 

It has been clear for some time, too, that among the Cephalopods, larvae and 
adults do not always have the same vertical distribution, so that an animal may be 
a temporary inhabitant of several distinct faunal zones during the course of its life 



history. Concerning vertical migration or diurnal rhythms in Cephalopods, we 
have no evidence, nor do we know how tolerant they are of changes in salinity. We 
do know, however, that some species are sensitive to changes in temperature and that 
a rapid lowering of temperature by two or three degrees is sufficient to kill some 
forms (e.g. Architeuthis and Sepia officinalis). 

The above is in broad outline a reasonable grouping of the species, but more 
exact limits may be possible when we know much more about this group. It must 
be admitted, for instance, that it is not easy to decide whether the distinction 
between a bathypelagic slope fauna and an oceanic bathypelagic fauna can be 

As the Madeiran Archipelago is the nearest oceanic group of islands to the Mediter- 
ranean, it may be pertinent to note that the following species in the list do not occur 
there : 

Slope animals. 
Spirula spirula. 
Lepidoteuthis grimaldi 
Cucioteuthis unguiculata. 
Enoploteuthis leptura. 
Tetronychoteuthis dussumieri. 
Architeuthis spp. 

Bathypelagic species. 
Mastigoteuthis schmidti. 
? Leachia cyclura. 
Taonius cymoctypus. 
Taonius pavo. 
Cranchia scabra. 
Japetella diaphana. 1 
Eledonella pygmaea. 
Vitreledonella richardi. 

Only two of the species, which may be termed slope and bathypelagic species, 
Histioteuthis bonelliana and Chiroteuthis veranyi, are found, and are known to breed, 
in the western Mediterranean. In this area the chief captures have been reported 
from places like Villefranche-sur-mer and Messina, that is, where upwelling from 
deep waters occurs. 

It is inappropriate here to discuss the cephalopods of the Mediterranean fauna at 
length, but it can be said that it is poor in bathypelagic species ; this is in agreement 
with what is known concerning the paucity of deep water species of other groups of 
animals (Ekman, 1953, p. 369). 


Alepisaurus ferox Lowe is well known as a voracious fish, and it has been known as 
a predator on squid since Verrill (1880, p. 234) described a brachial crown of Histio- 
teuthis from a fish captured off Nova Scotia in 1879. 

Examination of this series of 260 stomachs from Madeiran fishing grounds 
has yielded many comparatively rare species of squids which are not normally 
captured by plankton nets. The total of 18 species demonstrates that cephalopods 
can be regarded as regular items in the diet of this fish : 

1 " Dana " St. 1132, 36 , 10' N., 2° 46' W. in the Western Mediterranean near Gibraltar did yield 
one larval diaphana of 12 mm. (Thore, 1949, p. 28). We concur with Thore that this juvenile specimen 
was probably swept in through the Straits of Gibraltar in the easterly surface current. 



List of Species Eaten 

Number of 




number in 




one stomach. 



Heteroteuthis dispar . 





Loligo vulgaris . 





Abraliopsis morisii 





Pyroteuthis margaritifera 





Onychoteuthis banksi . 





Todarodes sagittatus . 





Ommastrephes pteropus 





Ommastrephids . 





Leachia cyclura 





Cranchia scabra 





Taonius pavo . 





Japetella diaphana 





Eledonella pygmaea . 





Octopus macropus 





Octopus sp. 





Alloposus mollis 





Ocythoe tuberculata 





Argonauta argo (see text) 


The dorsal mantle length has been taken as a standard measurement for size, 
and as a rough guide this is about half the total length of the animal in compact 
species like Heteroteuthis dispar, Abraliopsis morisii, Pyroteuthis margaritifera, 
Cranchia scabra, Japetella diaphana, Eledonella pygmaea, Alloposus and Ocythoe. 
It represents about only a third of that of Histioteuthis bonelliana, where the body 
is small and the arms disproportionately large. In the long and typical squids 
like Loligo, Onychoteuthis, Todarodes, Leachia and Taonius, mantle length is usually 
more than half the total length. 

Japetella diaphana, Alloposus mollis and Heteroteuthis dispar are the species most 
frequently represented and are perhaps rather sluggish creatures. It is particularly 
noticeable that among the swift-moving species the specimens of Onychoteuthis 
banksi were rather small, several being still in the planktonic stage. 

As will have been noted already, none of the cephalopods from the stomach of 
this fish can be regarded as truly deep-water species in the sense that this term can 
be applied to forms like V ampyroteuthis infernalis. All might reasonably be expected 
to occur in the upper 200 metres — the depth at which the fish were caught. 

Of the eighteen species, only four, namely Loligo vulgaris, Todarodes sagittatus, 
Japetella diaphana, Octopus macropus and Argonauta argo are common species in 
collections. Of the remainder there is little doubt that they are also common species, 
although records of capture are few. It is only recently that Japetella diaphana has 
been demonstrated to be one of the most abundant pelagic species of cephalopod 
(Thore, 1949), and Joubin (1933, p. 41) has similarly indicated from a preliminary 
examination of the " Dana " collections that the Cranchiidae (to which Leachia 


cyclura, Cranchia scabra and Taonius pavo belong) are also very numerous. Records 
of Ocythoe tuber culata (and possibly also Alloposus mollis to a lesser degree) are very 
few and do not imply scarcity but rather that the fishing gear hitherto employed by 
expeditions is unsuitable for capturing animals of this group. The same difficulty 
applies to the capture of swift surface species like Onychoteuthis banksi (Rees, 1949). 


Adam, W. 1941. Cephalopoda. Mem. Mus. roy. Hist. nat. (2) Fasc. 21 : 83-161, pi. i-iv and 

20 text-figs. 
— ■ — 1952. C6phalopodes. Res. Sci. Exped. Oceanog. Beige Eaux Coheres Africaines de 

I'Atlantique Sud, 3 (3) : 1, 142, pis. i-iii and 57 text-figs. 
Barnard, K. H. 1934. Cephalopoda from the Cape of Good Hope. /. Conch. London, 

20 : 44-45. 
Berry, S. S. 1916. The Octopod Ocythoe in California. Pomona J. Entom. Zool. 8 (1). 
1926. A note on the occurrence and habits of a luminous squid (Abralia veranyi) at 

Madeira. Biol. Bull. Woods Holl, 51 : 257-268, 2 text-figs. 
Bouxin, J., and Legendre, R. 1936. La fauna pelagique de I'Atlantique recueillie dans des 

estomacs de germons au large du golfe de Gascogne — Deuxieme Partie : Cephalopodes. 

Ann. Inst. Oceanog., Paris, 16 (1) : 1-99, 21 text-figs. 
Bruun, A. F. 1943. The biology of Spirula spirula (L.). " Dana " Rep. 24 : 1-46, 2 pis. 

and 13 text-figs. 
Chun, C. 1908. Ueber Cephalopoden der deutschen Tiefsee — Expedition. Zool. Anz. 33 : 

1910. Die Cephalopoden. I : Oegopsida. Wiss. Ergeb. Tiefsee-Exped." Valdivia 18 : 

Crosse, H., and Fischer, P. 1862. Nouveaux documents sur les Cephalopodes gigantesques. 

/. de Conch. Paris (3), 2 : 124-140. 
Degner, E. 1925. Cephalopoda. Danish Oceanog. Exped. 1908-10, 11, C.l : 1-94, 52 text- 
figs, and 3 charts. 
Drouet, H. 1858. Mollusques marins des lies Acores. Mem. Soc. Agric. Sci., Belle Lettres, 

Aude (II), 2. 
Ekman, S. 1953. Zoogeography of the Sea. London: Sidgwick & Jackson, xiv -f 417 pp. 
Ferussac, A. de, and Orbigny, A. d'. 1835-1848. Histoire naturelle generate et particuliere 

des Cephalopodes acetabuliferes vivants et fossiles. Paris. 2 vols., text and atlas. 
Fischer, H., and Joubin, L. 1906. Note sur les Cephalopodes captures au cours des expedi- 
tions du " Travailleur " et du " Talisman." Bull. Mus. Hist. Nat. Paris, 1906 : 202-205. 

1907. Cephalopodes. Exped. sci. " Travailleur " et du " Talisman," 8 : 313-353. 

Girard, A. A. 1892. Les Cephalopodes des iles Acores et de l'ile de Madere. /. Cien 

Math. Fis. Na. Lisboa, 1892. 
Harvey, E. N. 1952. Bioluminescence. New York : Academic Press, xvi and 649 pp., 

187 text-figs. 
Hertling, H. 1938, Ueber eine auf Juist gestrandete Sthenoteuthis caroli, (Furtado). Wiss. 

Meeresuntersuch. 1 : 93-111. 
Hoyle, W. E. 1884. On Loligopsis and some other genera. Proc. roy. Phys. Soc. Edin. 

8 : 313-333- 
1885. Report on the Cephalopoda. Rep. Sci. Res. H.M.S. " Challenger," Zool. 16 (44 : 

1-245, pis. i-xxxiii and 10 text-figs.). 
Joubin, L. 1895. Contribution a l'etude des Cephalopodes de I'Atlantique Nord. Res. 

Camp. Sci. Monaco, 9. 
1900. Cephalopodes provenant des campagnes de la " Princesse Alice " (1891-1897). 

Ibid. 17. 


1920. Cephalopodes provenant des campagnes de la " Princesse Alice " (1898-1910). 

Ibid. 54 : 1-95. 16 pis. 

1924. Contribution a l'6tude des Cephalopodes de l'Atlantique Nord. Ibid. 67 : 1-113. 

14 pi. and 6 tables. 

1933. Notes preliminaries sur les Cephalopodes des Croisieres du Dana (1921-1922). 

4 C Partie. Ann. Inst. Oceanog., Paris, 13 (1) : 1-49, 48 text-figs. 

Lesueur, C. A. 182 1. Descriptions of several new species of Cuttlefish. /. Acad. Nat. Sci. 

Philad. 2 (1) : 86-101, 4 pi. 
Lonnberg, E. 1896. Notes on some rare Cephalopods. K. Vetensk-Akad. Forhandl. 1896, 

No. 8 : 603-612, 4 text-figs. 
Mac Andrew, R. 1852. Note of the mollusca observed during a short visit to the Canary and 

Madeira Islands, etc., in the months of April and May, 1852. Ann. Mag. Nat. Hist. (2), 

10 : 100-108. 
Nobre, A. 1937. Moluscos testaceos marinhos do Arquipelago da Madeira. Mem. Mus. 

Zool. Univ Coimbra, 1937 ( x ) No. 98 : 1-101. 
Odhner, N. H. 193 i. Beitrage zur Malakozoologie der Kanarischen Inseln. Lamelli- 

branchien, Cephalopoden, Gastropoden, Arch. Zool. Stockholm, 23A (14) : 1-116, 2 pis. 

and 48 text-figs. 
Pfeffer, G. 191 2. Die Cephalopoden der Plankton Expedition. Ergeb. Plankton-Exped. 

Humboldt-Stiftung 2, F,a : 1-8 15, text and atlas. 
Rees, W. J. 1949. Note on the hooked squid, Onychoteuthis banski. Proc. Malac. Soc. 

London, 28 : 43-45. 

1950. On a giant squid, Ommastrephes caroli, Furtado, stranded at Looe, Cornwall. 

Bull. Brit. Mus. Zool. 1 (2) : 31-41, 2 pis., 6 text-figs, and 3 maps. 

Robson, G. C. 1929-1932. A Monograph of the Recent Cephalopoda, Parts I and II. London : 

British Museum (N.H.). 
Salisbury, A. E. 1953. Mollusca of the University of Oxford Expedition to the Cayman 

Islands in 1938. Proc. Malac. Soc. 30 (1 and 2) : 39-54, pis. 7 and 8. 
Sasaki, M. 1929. A Monograph of the dibranchiate cephalopods of the Japanese and adjacent 

waters. /. Fac. Agric. Hokkaido Imp. Univ., Sapporo, 20, Suppl. No. 1-357, 3° pl s - an( i 

159 text-figs. 
Sverdrup, H. U., Johnson, M., and Fleming, R. 1946. The Oceans, their Physics, Chemistry 

and General Biology. New York. 
Thore, S. 1949. Investigations of the " Dana " Octopoda : Part I : Bolitaenidae, Amphi- 

tretidae, Vitreledonellidae and Alloposidae. " Dana " Rep. 33 : 1-85, 69 text-figs. 
Verrill, A. E. 1879-80. The cephalopods of the north-eastern coast of America. Part I : 

The gigantic squids (Architeuthis) and their allies, with observations on similar large 

species from foreign localities. Trans. Conn. Acad. Sci. 5 : 177-257 ; pis. 13-25. 

1880-81. The cephalopods of the north-eastern coast of America. Part III : the 

smaller cephalopods including the " squids " and the octopi with other allied forms. Ibid. 
5 : 259-446, pis 26-56 and 3 text-figs. 

Watson, R. B. 1897. On the marine mollusca of Madeira ; with descriptions of thirty-five 
new species, and an index-list of all the known sea-dwelling species of that island. /. 
Linn. Soc. 26 : 233-329, pis. 19 and 20. 



By W. J. REES, D.Sc. 

(British Museum (Natural History)) 




Introduction .......... 283 


Eledone cirrhosa (Lamarck) 

(a) Spawning 

(b) The Egg Masses 

(c) Description of the larvae 



Eledone moschata (Lamarck) 

(a) Egg Masses and early stages 



Distribution of the European Species 



The Differences between E. cirrhosa and E. 




References ...... 





The newly hatched larvae of the Lesser Octopus, Eledone cirrhosa (Lamarck), are described 
and figured for the first time from material collected by the Fishery Board for Scotland. 
Records of eggs and spawning have been brought together and additional ones listed. The 
eggs and larvae of this species are compared with those of the Mediterranean E. moschata and 
the distribution of both species reviewed. E. moschata occurs throughout the Mediterranean 
and its distribution outside is limited to neighbouring coasts, north and south of the Straits of 
Gibraltar. The records show that E. cirrhosa occurs in the western Mediterranean and along 
the western coasts of Europe to Iceland, the Faroes and northwards to Trondhjem on the 
Norwegian coast. The characteristics of the two species are compared. 


In this paper some new observations are made on the eggs and larvae of the Lesser 
Octopus (Eledone cirrhosa Lamarck) and our previous knowledge of its reproduction 
and distribution is summarized. Early naturalists confused this species with Eledone 
moschata (Lamarck) and it has been deemed desirable to bring together what is known 
of the eggs and larvae, as well as the distribution of both species to enable a summary 
of the differences between the species to be presented. 1 

Although Eledone cirrhosa is much more widely distributed in North European 
waters than the Common Octopus (Octopus vulgaris Lamarck), it is surprising that 
so little is known about its habits and life history. Its morphology and anatomy was 
the subject of a memoir by Isgrove (1909). 

1 The larva of a South African Eledonid, Pareledone nigra, has recently been described by me (see Rees, 



Text-fig. i. — The recorded distribution of Eledone cirrhosa in inshore waters. Trawling 
records from the central North Sea are not included. The sources of all the records are 
given in the bibliography. 



(a) Spawning 

The spawning of Eledone cirrhosa in captivity was first noted by Joubin (1888) 
who observed it in an aquarium at Banyuls in the month of June. According to 
Joubin there were about 30 groups of eggs (and traces of another 30) most of them 
being eaten by the female. Each bunch contained 5-19 eggs, the greatest number 
being laid first. The eggs were white in colour, each being 7-8 mm. in length. 

Spawning of Eledone was also noted by Gravely (1908) in an aquarium tank at 
Port Erin, Isle of Man, in July. He noted that eggs were about 7 mm. by 2-5 mm. in 
diameter and that one to four bunches of eggs were laid almost every day for about 
a month, after which spawning was less regular and was soon over. 

Isgrove (1909) states that one Eledone lays about 800 eggs and that these are 
spawned in groups of 25-30 eggs. Egg clusters 4-7 cm. long were collected in Aberdeen 
market by Bowman ; the full sized ova were 8-9 mm. in length and about 4 ■ o mm. 
in diameter at the broad end (Russell, 1922). It now seems that these eggs are a little 
too large to be those of E. cirrhosa, but their identity cannot be known for certain 
until we have some information on the eggs of Graneledone verrucosa (Verrill) ; this 
species replaces E. cirrhosa to the north of the Faroes. 

Spawning in aquaria has been noted at Plymouth in January (Marine Biological 
Association, 1931) and in July (Isgrove, 1909) ; at Port Erin in July (as noted by 
Gravely) and in September (Moore, 1937, p. 196). A female captured on the Dogger 
Bank spawned in the Heligoland aquarium in January (Hertling, 1936, p. 294) 
Stephen (1944, p. 252) mentions several clusters of ova from N.N.W. of Ronas Voe, 
Shetland (position, 6o° 42' N., i° 46' W.) trawled on 4th April, 1927, and, as the 
embryos were well developed it can be assumed that spawning occurred in February 
or early March. 

From this it appears that Eledone may spawn all the year round, and this is 
borne out by records of larvae, noted by Stephen (1944, p. 251). He observed that 
although larvae were present all the year in the plankton catches of the Fishery Board 
for Scotland, they were more frequent during the period May to August ; this 
suggests maximum spawning in April, May, June and July. As to whether this 
period of more intensive spawning is linked with the known seasonal migration of 
Eledone into inshore waters we have insufficient evidence. Even the kind of haunt 
chosen for spawning in nature is not known for certain, but it appears that Eledone 
does not brood over its eggs, nor does it seem to lay them in shells or pots, as does 
Octopus vulgaris, for it has never been taken with its eggs. 

(b) Egg masses 

Apart from eggs seen in aquaria, Eledone spawn is rarely taken and only two 
positive records are known to me, the batches trawled near Ronas Voe in the Shet- 
lands (Stephen, 1944) and a very large cluster from the Eddystone Grounds off 
Plymouth in the collections of the Plymouth Laboratory. 

The large egg mass from the Eddystone Grounds contains a very large number of 
undeveloped eggs. The stalks of the eggs are very short and the largest eggs are 67 


mm. in length by 2-4 mm. in width. Some eggs are extremely small, being only 
2-85-3-6 mm. in length by 1-0-1-5 mm. in diameter (PL 9). 

The small egg-cluster from Ronas Voe contains embryos in an advanced stage of 
development. The eggs themselves are 6-65-6-79 mm. in length by 2-94-3-29 mm. 
in width. There is still a large yolk mass, but the embryos are well formed. Chroma- 
tophores are developing the arms, head and body and the Kollikersche buschel are 
clearly seen on the head and mantle (PI. 10, figs. 1-4). At this size (3 mm. in. ventral 
mantle length) the single row of suckers on the subequal arms are formed. . 

A third cluster in the British Museum is without any particulars ; the eggs are 
poorly preserved and resemble those from Ronas Voe. 

Records of larvae of E. cirrhosa are few. Lo Bianco (1909) found young Eledone 
in the plankton in the Bay of Naples ; those found in April had a length (? 
total length) of 40 mm. and those found in October a length of 120 mm. He gives no 
adequate description and there is no certainty as to which species he had. Scottish 
records of larvae are given by Russell (1922) and Stephen (1944). Russell noted that 
the arms are much shorter in proportion to the body (3 : 5) in the young, the back is 
smooth and covered with large chromatophores and the body is generally surrounded 
by a thick, soft, translucent cuticle. 

In the Bay of Biscay area Bouxin and Legendre (1936, p. 24) found seven specimens 
ranging from 21-33 mm. in length in the stomachs of germon in positions approxi- 
mately 90-100 miles to the south-west of Glenans and at about 250 miles to the 
south-west (that is, near Cape Finisterre). 

A new description is therefore needed and is given below. 

(c) Description of the larvae 

Dr. A. C. Stephen has kindly allowed me to examine a series of 18 larvae, from the 
catches of the Fishery Board for Scotland, which were reported by him in 1944. 

It has already been noted that larvae still in the eggs may reach a ventral mantle 
length of 3 mm. in large eggs. In small eggs this length would presumably be less, 
so that planktonic larvae of approximately this size can be regarded as having been 
taken within a few days of hatching. 

The young larva (PL 10, figs. 5-6) has much the same shape as the larva of 
Octopus vulgaris (the only species it could be confused with in British waters) with its 
short arms and rather squat mantle. In details, however, there are big differences, 
the most noticeable feature being the arrangement of the chromatophores on the 
mantle. Here the entire mantle is uniformly covered with large reddish brown 
chromatophores, and these are also prominent on the head and arms. On the latter 
there is a single row and those of the central portion of the head are deep-seated. 
Overlying these in the transparent outer integument are other fainter reddish brown 
chromatophores, which increase in number and size with growth. The outer integu- 
ment of the eye is silvery and sometimes has a greenish hue. 1 

The arms are subequal and do not have the thin cirriform tips seen in very young 
larvae of Octopus vulgaris. No. 12 (Table I) has 6-7 fully formed suckers on each arm 

1 All references to colour mean colour of preserved specimens in alcohol. 



with the clear rudiments of 8-9 more at the tip of the arm. A slightly larger specimen 
(No. 13) has 9 suckers on each arm and about 8 rudiments at the tip. 

The largest larva available (No. 19) has about 28 suckers plus rudiments at the tip. 
The first proximal sucker has a diameter of 0-56, the second, third and fourth have a 
diameter of 07, 0-84 and 1-05 respectively. Suckers 5-7 are the largest and then they 
diminish in size distally. The web is subequal reaching to the 7th-8th suckers, that is, 
to about one-third of the length of the tentacles. This specimen taken on 27th 
November, 1930, was recorded from square E 13 b by Stephen (1944). 

This late larva has developed a mantle fin-ridge as in the adult. The ventral 
mantle is smooth but the dorsal mantle and head are both covered with tubercles. 
The larval chromatophores are still discernible because of their large size, but the 
areas in between have become covered with a large number of small chromatophores. 

As noted above, the arms of the larval Eledone are quite short at hatching from 
the egg and they are usually not so long as the ventral mantle (Table I). Growth of 
the arms of the planktonic larvae is more rapid than mantle length (Figs. 3 and 4) 
and is also reflected in the relation of mantle length to total length (Fig. 2) . 

Table I. 

— Larvae Collected by the Fishery Board J 

or Scotland 













Total length . 











Dorsal mantle length 



eye) . 











Ventral mantle length 

3-6 5 





3'8 5 





Head width . 











Mantle width . 











Diameter of eye 





1 -i 

1 -2 

I -2 

1 -26 



Length of arms 











Diameter of suckers 
























Total length . 










Dorsal mantle length 



eye) . 










Ventral mantle length 



4 - 4 I 







Head width 



3 '99 







Mantle width . 

■ 3-8 5 









Diameter of eye 

1 -26 

1 -26 

i- 4 







Length of arms 

■ 3*22 









Diameter of suckers 

• o-3 








I -12- 

I -2 


(a) Egg masses and larvae 

We know less about the spawning and larval stages of this species than we do of 
Eledone cirrhosa. Korschelt (1893) described an egg-mass found on a Pinna shell at 


— ~T T 

i ■ ■ 

i i i i 

i • i i i i 1 r 1 1 1 i 

i i i r— t — 





1 1 ' T 1 





. Z 












i i 


1 1 1 1 

i i i i i i i i i i i i 

1 1 1 1 1 


10 15 20 25 30 

Text-fig. 2. — The relation of mantle length to total length in larvae of 
Eledone cirrhosa. Measurements are in mm. 



-| r — t - - » 1 — « I — '■ " ■ | — — r 1 « 1 1 1 « 1 ~» 1— — i T 


h < 

op o 

o ^o 

°8> °° 


J •— i ■ i t i i \ i i i l i i i i * u 

2 4 6 8 10 12 

Text-fig. 3. — The relation of arm length to ventral mantle length in larvae of 
Eledone cirrhosa. Measurements are in mm. 


Rovigno in the Adriatic. He gave the total number of eggs as 65-70 ; this mass 
was made up of small clusters of 2-4 eggs joined to the main mass by a common 
stalk. The eggs (excluding stalk) were 15 mm. in length, that is, the same length as 
the eggs figured by Jatta (1896, tav 7, fig. 3). 

Korschelt was under the impression that he had the eggs of E. aldrovandi (i.e., 
cirrhosa) , but this mistaken view was corrected with the identification of the large 
eggs with E. moschata by Jatta and confirmation by Gravely (1908) that the smaller 
eggs belonged to E. cirrhosa. There have been embryological studies on development 

°"T T" 1 J 1 — T 1 














O O 





2 4 6 8 10 12 

Text-fig. 4. — The relation of sucker diameter to ventral mantle length in larvae 
of Eledone cirrhosa. Measurements are in mm. 

in the egg (Sacarrao, 1943, 1945, 1951 and 1952) but there is no detailed description 
of the newly hatched larva. Sketches of juvenile Eledone have been published by 
Jatta (1896, tav 7, figs. 5 and 10), while Naef (1923) gives a drawing of a post-larval 
E. moschata from Trieste. A new description is much needed. 


Eledone cirrhosa (Lamarck) is found in the Mediterranean, but its full distribution 
there is not known. It appears to be a common species in the western part, but there 
are no records for the eastern Mediterranean. Under the name Eledone aldrovandi 
there are numerous records for the western Mediterranean (Jatta, 1896), but there 
are no records east of Dalmatia (Robson, 1932, p. 266) and according to Ninni 
(1884) it does not penetrate to the head of the Adriatic. 

In the Atlantic E. cirrhosa has a much less restricted distribution than E. moschata. 

zool. 3, 6. 21 


It is a common species on the European continental shelf extending to southern 
Iceland, the Faroes (Brunn, 1945, p. 8) and the west coast of Norway. In the open 
areas of the shelf it appears not to extend beyond the line Iceland-Faroes-Bergen, but 
is a common species southwards of this line. On the Norwegian coast itself it reaches 
Ostnesfjord in the Lofotens, but is said to be scarce north of the Trondhjemsfjord 
(Grieg, 1933). 

Its distribution in inshore British waters as recorded in the literature on cephalo- 
pods is plotted on Map I, and, quite apart from the fact that such maps tend to 
reflect areas worked by biologists, the species appears to be scarcer in the southern 
North Sea than elsewhere. Records from trawling grounds offshore have not as a rule 
been plotted (those from the North Sea trawling grounds are given by Grimpe, 1925), 
and it has not been possible to search for all the occasional records buried in the 
transactions of local natural history societies. 

Eledone moschata (Lamarck) is a Mediterranean species which ranges from Istanbul 
(Digby, 1949), the Syrian coast (Gruvel, 1931) and Palestine (Bodenheimer, 1937) to 
the adjoining region of the Atlantic. Korschelt (1893, p. 68) implies that it is the 
common Eledone of the Adriatic and it has also been found commonly at Naples 
(Jatta, 1896, and Naef, 1923), the Gulf of Marseilles (Vayssiere, 1917) and Monaco 
(Boone, 1933) to quote only a few of the more recent records. Outside the Mediter- 
ranean the species appears to be rare and records are few. The " Talisman " took a 
female specimen in the Bay of Cadiz from a depth of 60 m. (Fischer & Joubin 
1907, p. 328), while Adam (1941, p. 140) reported the first specimen from the African 
Coast (Baie du Levrier, Port-E'tienne, Mauritania). 

There are no authentic records from northern Europe ; Nielsen's Eledone moschata 
from the Faroes (1930) proved on re-examination to be E. cirrhosa (Brunn, 1945, p. 9). 


The main differences between the species are tabulated below. 

Eledone cirrhosa Eledone moschata 

No musk odour. . Musk odour. 

Skin with warts on dorsum. . Skin smooth. 

Ridge along edge of mantle. . No pallial ridge. 

The seven non-hectocotylized arms of the . The seven non-hectocotylized arms carry a 

male carry close-pressed, flattened suckers, double series of transverse lamellae at their 

forming cirri at the tips of the arms. tips. 

The two retractor muscles of the gills are . The two retractor muscles of the gills are 

fused at their base and form a " Y " inserted separately. 

Spermatophore with spines. 1 . Spermatophore without spines. 

Colour of adult : Light yellowish brown with . Colour of adult : Greyish brown colour with 

diffuse rust-brown patches on the back. darker almost black patches on the dorsal 

Ventral mantle pale ivory or pinkish side. Preserved animal is grey to dark 

yellow with a greenish iridescent sheen. grey with numerous dark patches. 

Eggs, Pyriform, ca. 7 x 2-5 mm. in clusters . Eggs sausage-shaped ca. 15 x 4 mm. in 

of about 30. clusters of two to four. 

1 Fort (1941) creates a new genus Acantheledone for E. cirrhosa laying particular stress on this character. 


Eledone cirrhosa (cont.) Eledone moschata (cont.) 

Newly hatched young ca. 3-0 mm. in ventral . Newly hatched young probably ca. 8 mm. 

mantle length. in ventral mantle length. 

Parasites : . Parasites : 

Dicyemennea eledones (Wagner, 1857). D. eledones. 

Chromidina coronata (Foettinger, 1881). D. moschatum Whitman, 1882. 

Adam, W. 1933. Notes sur les cephalopodes. Ill : Les Cephalopodes du Sud de la mer du 
Nord. Bull. Mus. Roy. Hist. Nat. Belg. 9, No. 46 : 1-45 Charts A-D. 

1939. Notes sur les cephalopodes. X : Sur quelques cephalopodes de la mer d'Islande. 

Ibid. 15, No. 16 : 1-13, 4 text-figs. 

1 94 1. Resultats scientifiques des croisieres du Navire — licole Beige Merator. 

Cephalopoda. Mem. Mus. roy. Hist. Nat. Belg. (2), Fasc. 21 : 83-162, pi. i-iv and 20 text-figs. 
Bodenheimer, F. S. 1937- Prodromus faunae Palestinae. Mem. Inst. Egypte, 33 : 1-286, 

4 text-figs. 
Boone, L. 1933. Scientific results of cruises of the yachts " Eagle " and " Ara " 1921-1928, 

William K. Vanderbilt, Commanding. Coelenterata, Echinodermata and Mollusca. Bull. 

Vanderbilt Mus. 4 : 1-2 17. 
Bouxin, J., and Legendre, R. 1936. La faune pelagique de l'Atlantique recueillie dans les 

estomacs de germons au large du golfe de Gascogne. Deuxieme Partie : Cephalopodes. 

Ann. Inst. Oceanogr. Paris 16 (1) : 1-99, 21 text-figs. 
Bruun, A. F. 1945. Cephalopoda. Zoology of Iceland, 64 (64) : 1-15, 4 text-figs. 
Chumley, J. 1918. The Fauna of the Clyde Sea Area. Glasgow. 
Clarke, W. J. 1936. Yorkshire Cephalopoda. /. Conch. Lond. 20 (9) : 257. 
Dautzenberg, P., and Fischer, P. H. 1925. Les mollusques marins du Finistere et en 

particulier de la region de Roscoff. Trav. Stat. Biol. Roscoff, Fasc. 3 : 1-180. 
Degner, E. 1925. Cephalopoda. Danish Oceanogr. Exped., 1908-10, II, C. 1 : 1-94, 52 

Digby, B. 1949. Cephalopods from local waters at the University of Istanbul. Nature, 163, 

No. 4141, p. 411. 
Evans, W. 1899. Notes on the effects of the recent October gale upon marine life on the 

coasts of the Lothians. Ann. Scott. Nat. Hist. 29 : 6-1 1. 
Fischer, P. 1867. Catalogue des Nudibranches et Cephalopodes des cotes oceaniques de la 

France. /. de. Conch., Paris, 15 : 1-15. 

and Joubin, L. 1907. Cephalopodes. Exped. Sci. " Travailleur " et du " Talisman." 

8 : 313-353- 
Forbes, E., and Hanley, S. 1853. A History of British Mollusca. London : J. van Voorst. 

Vol. IV. 
Forrest, J. E., and Waterston, A. R. 1934. Scottish records of Cephalopods. I : A 

" Cephalopod Year " in the Clyde. Scot. Nat. 1934 : 29. 
Fort, G. 1937. Note sur les especes francaises du genre Eledone. Bull. Soc. Hist. Nat. 

Toulouse, 71 (3) : 309-314. 

1 94 1. Eledone Leach et Acantheledone n. g. deux genres fondes sur la structure des sperma- 

tophores (Moll. C6phalopodes) C.R. Acad. Sci. Paris, 212 (17) : 724-726. 

Godfrey, R. 1900. Eledone cirrosa in West Lothian. Ann. Scot. Nat. Hist. 1900 : 185. 
Gravely, F. H. 1908. Notes on the spawning of Eledone and on the occurrence of Eledone 

with suckers in double rows. Mem. Manchester Lit. Phil.-Soc. 53 (4) : 1-14, 2 pis. 
Grieg, J. A. 1933. Cephalopods from the West Coast of Norway. Bergens. Mus. Aarb. 1933, 

No. 4 : 1-25, pi. i-iv and 1 text-fig. 
Grimpe, G. 1925. Zur Kenntnis der Cephalopoden fauna der Nordsee. Wiss. Meeresunter- 

suchungen. Biol Anst. Helgoland, 16 (3) : 1-124, 1 pi. and 34 text-figs. 


Gruvel, A. 193 i. Les Etats de Syrie. Richesses marines et fiuviales. Exploitation actuelle, 

453 PP-> 30 Pis. 
Gyngell, W. 1928. Cephalopoda landed at Scarborough in 1927. /. Conch. Lond., 18 : 216. 

1929. Cephalopoda etc. at Scarborough in 1928. /. Conch., 18 (10) : 287. 

Haddon, A. C. 1886. Recent Contributions to the Marine Invertebrate Fauna of Ireland. 

Zoologist (3) 10 : 1-7. 
Haines, F. H.,1945. Notes on general zoology, 1944. Mollusca. Pap. Hampshire Field Club, 

16 (2) : 233. 
Hargreaves, J. A. 1910. Marine Mollusca of the Yorkshire Coast. /. Conch. Lond. 13 (4) : 

Hertling, H. 1936. Mitteilungen iiber Todaropsis eblanae (Ball), Octopus vulgaris L. und 

Eledone cirrosa (Lam.) aus. der Nordsee. Zool. Anz. 114 (11-12) : 289-296, 1 text-fig. 
Horsman, E. Additions to the marine fauna of the Aberystwyth district. Aberystwyth Studies, 

4 : 259-269. 
Hoyle, W. E. Report on the Cephalopoda. Rep. Sci. Res. H.M.S. " Challenger," Zool. 16 

(44) : 1-245 pis. 1-33, 10 text-figs. 
Isgrove, A. 1909. Liverpool Mar. Biol Comm. Memoir XVIII : Eledone. 
Jatta, G. 1896. I cephalopodi viventi nel Golfo di Napoli. Fauna und Flora des Golfes von 

Neapel. Monogr. 23 : 1-268, pi. 1-3 1 and 64 text-figures. 
Jeffreys, J. G. 1869. British Conchology , London : J. van Voorst. Vol. V. 
Joubin, L. 1888. Sur la ponte de YEledone et de la seiche. Arch. zool. exper. et gen. (2) 

6 : 155-163, 1 text-fig. 

1920. Cephalopodes provenant des Campagnes de la " Princess Alice " (1898-19 10) 

3 e S6rie). Res. Camp. Sci. Monaco, 54 : 1-95, 16 pi. 

Korschelt, E. 1893. Uber den Laich und die Embroynen von Eledone. Sitz. Ber. Get. 

Naturf. Berlin, 1893, No. 2 : 68-73, 2 text-figures. 
Lo Bianco, S. 1909. Notize biologiche riguardante specialmente il periods maturita sessuale 

degli animali del Golfo di Napoli. Mitth. Zool. stat. Neapel, 19 : 513-761. 
Marine Biological Association, 1931. Plymouth Marine Fauna. Second Edition. 
Massy, A. L. 1909. The Cephalopoda Dibranchiata of the Coasts of Ireland. Fisheries Ireland 

Sci. Invest. 1907, I (1909) : 1-39, PI. I— III. 
1 91 3- Further records of the Cephalopoda Dibranchiata of the Coast of Ireland. 

Ibid. 1912, V (1913). 
1928. The Cephalopoda of the Irish Coast. Proc. Roy. Irish Acad. 33 {Section B), 

(2) : 25-37. 
Moore, H. B. 1937. Marine Fauna of the Isle of Man. Proc. and Trans. Liverpool Biol. 

Soc. 50 : 1-293. 
Naef, A. 1921-28. Fauna e Flora del Golfo di Napoli. Monogr. 35 : Die Cephalopoden. 
Nichols, A. R. 1900. A list of the Marine Mollusca of Ireland. Proc. Roy. Irish Acad. 5 

(4) : 477-662. 
Nielsen, E. 1930. Cephalopoda. Zoology of the Faroes. Part 56. 
Ninni, A. 1884. Catalogo dei Cefalopodi Dibranchis osservati nell'Adriatico. Att. Acc.Ven.- 

Tren.-Instr. Padova, 9 : 159. 
Nordga ard , . 1 92 3 . The Cephalopoda Dibranchiata observed outside and in the Trondh j em- 
fjord. Kgl. Norske. Vidensk. Selsk. Shrift. 1922, No. 5 : 1-14. 4 text-figs. 
Rees, W. J. 1954. The larva of Pareledone nigra (Hoyle) from South Africa. Proc. malac. 

Soc. 31 : 50-51, 1 pi. 
Rendall, R. 1931. Notes on Eledone cirrosa Lam. /. Conch. Lond. 19 : 161-162. 
Robson, G. C. 1932. A Monograph of the Recent Cephalopoda. Part II : The Octopoda. 

British Museum (Nat. Hist.), London. 
Russell, E. S. 1922. Report on the Cephalopoda collected by the research steamer " Gold- 
seeker " during the years 1903-1908. Fisheries Scotland Sci. Invest. 1921, III : 1-45, pis. 



Sacarrao, G. F. 1943. Observations sur les derniers phases de la vie embryonnaire de 
" YEledone." Arq. Mus. Bocage Lisboa, 14 : 25-35, 1 pi. and 3 text-figs. 

1945- Etudes embryologiques sur les Cephalopodes. Ibid. 16 : 33-68, 9 text-figs. 

1951. Notice on the embryonic shell sac of Octopus and Eledone. Ibid. 22 : 103-105, 

PI. I. 

1952. Quelques remarques sur une comparison des ontogeneses d'Octopus et d'Eledone. 

Rev. Fac. Cienc. Lisboa (2) C, 2 (2) : 215-222. 

Service, R. 1892. Eledone cirrosa Lamarck in the Solway Firth. Ann. Scott. Nat. Hist. 

1892 : 202. 
Stephen, A. C. 1934. Scottish records of Cephalopods. Scot. Nat. 1934 : 29. 
— ■ — ■ 1944. The Cephalopoda of Scottish and adjacent waters. Trans. Roy. Soc. Edin. 61 

(1), No. 9 : 247-270, 14 text-figs. 
Stevenson, J. A. 1935. The Cephalopods of the Yorkshire Coast. /. Conch., London, 

20 (4) : 102-116. 
Thorson, G. 1946. Reproduction and larval development of Danish marine bottom inverte- 
brates, with special reference to the planktonic larvae in the Sound (0resund). Medd. 

Komm. Dansk. Fisk. — og Havundersog. Ser. Plankton, 4 (1) : 1-523, 199 text-figs. 
Tomlin, J. R. le B. 1892. Notes on the Marine Mollusca of the North Wales coast with 

complete lists of the recorded nudibranchs and cephalopods. /. Conch. Lond. 7 (1) : 25-31. 
Tregelles, G. F. 1896. The Marine Mollusca of Cornwall. Ibid. 8 (6) : 191-200. 
Vayssiere, A. Note sur la presence d'un bras supplementaire chez un Eledone moschata 

Leach. /. de Conch. Paris, 63 : 123-125, 1 pi. 


" a 1956 


Egg mass of Eledone cirrhosa (Lamarck) from the Eddystone Grounds, Plymouth. 

■nil. B.M. {N.H.) Zoology 3, 6 


Egg mass of Eledone cirrhosa (Lamarck) 

Eggs and larvae of Eledone civvhosa (Lamarck) del. G. L. Wilkins. 

Fig. i. — Portion of an egg cluster from Ronas Voe, Shetlands. 

Fig. 2. — A well developed larva in situ. 

Fig. 3-4. — Views of larvae removed from their egg capsules. 

Fig. 5-6. — Side and ventral views of a planktonic larva from the catches of the Fishery 
Board for Scotland. 

hill. B.M. (.V.//.) Zoology 3, 6 





I f 

«i: " -; • 

EM- • .'-S 

fey ' ■•;■' 

Eggs and larvae oi Eledone cirrhosa (Lamarck) 

ENT -Q^ 

?&£ <f * 


- 2 MAR 1956 







ZOOLOGY Vol. 3 No. 7 

LONDON: 1956 




(East African Fisheries Research Organization, Jinja, Uganda) 

Pp. 295-333 ; 10 Text-figures. 



ZOOLOGY Vol. 3 No. 7 

LONDON: 1956 

(NATURAL HISTORY), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical Series. 

Parts will appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

This paper is Vol. 3, No. 7 of the Zoological series. 


Issued February, 1956 Price Ten Shillings 





Synopsis of Genera 

Genus Macropleuro 
Generic charac 

Macropleuro dus bic 
Syncranium ai 
Coloration anc 
Study materia 

Genus Platytaeniod 
Generic charac 

Platytaeniodus dege 
Study materia 

Genus Hoplotilapia 
Generic charac 

Hoplotilapia retrode, 
Study materia 

Genus Paralahidoch 

Paralabidochromis i 
Summary . 

of tb 

:ters a 

olor (] 

id ass 

I and 
ters a 

ni Bo 

I and 
ters a 
ns Hi 




e Haplochromis Grou 

nd synonymy 
3oulenger) . 

ociated musculature 

distribution records 
nd synonymy 

distribution records 
nd synonymy 

distribution records 
nov. . 

%e sp. nov. 

p Occ 









ZOOL. 3, 



In his revision of the Lake Victoria Cichlidae, Regan (1922) recognized four endemic 
monotypic genera, the species being Astatoreochromis alluaudi Pellegrin, 1904, 
Hoplotilapia retrodens Hilgendorf, 1888, Platytaeniodus degeni Boulenger, 1906, and 
Macropleurodus bicolor (Boulenger), 1906. Subsequent collections made by the 
Cambridge Expedition (1930-1931) extended the range of A. alluaudi to include 
the Lake Edward system and Lakes Kachira and Nakavali (Trewavas, 1933), but 
provided no further distributional data for the other three genera. Since Astatoreo- 
chromis alluaudi occurs beyond the Victoria system, it is preferable to delay revision 
of this species until numerous specimens from the different localities can be examined. 
Some notes on the osteology and possible phyletic relationships of Astatoreochromis 
have been published already (Greenwood, 1954). 

A fifth genus which, but for its geographical separation, would not have been 
distinguished from Labidochromis of Lake Nyasa, is recorded for the first time. 

The present study is based on specimens collected by and for the East African 
Fisheries Research Organization during a field study of the Lake Victoria Cichlidae, 
and on material in the collections of the British Museum (Nat. Hist.), Museum 
National d'Histoire naturelle, Paris, and the Museo Civico di Storia Naturale, 
Genoa. It forms the first part of a revision of the Lake Victoria Haplochromis 
species flock. 

Notes on counts and measurements 

The counts and measurements used are as defined by Trewavas (1935), except that 
" length of head " is measured directly from the posterior margin of the operculum 
to the premaxillary symphysis. This method has been found to yield more consistent 
results than measurements taken between verticals through the posterior tip of the 
operculum and the level of the tip of the snout, along a line parallel to the longitudinal 
axis of the body. Likewise, length of snout is measured directly. 
Other measurements and counts are : 

Depth of preorbital : measured from about the middle of the orbital rim of the 
preorbital bone along a line which continues the radius of the eye at this point ; the 
line approximately bisects the bone. 

Interorbital width : the least width of the roofing part of the frontal bones, 
between the eyes. 

Lower jaw : measured directly from the angle to the symphysis. 

Lateral line scale series : After last upper lateral line scale, proceed to the scale 
of the lower lateral line next behind the transverse row that includes the last scale 
of the upper lateral line and slopes downwards and forwards from it. 

In addition to those defined by Trewavas. Depth of cheek : the greatest depth 
measured vertically from the lower orbital margin to the lower edge of the adductor 
mandibulae muscles. In most specimens this is virtually a measurement of the depth 
of the scaled portion of the cheek. 


Post-ocular part of the head : measured directly from the posterior orbital margin 
to the posterior tip of the operculum. 


1. Anal spines three ............ 2. 

Anal spines four or more ........ Astatoreochromis 

2. Teeth of upper jaw in two or more series anteriorly, but only in a single (rarely 

double) series laterally ........... 3. 

Teeth of upper jaw in 2-5 series both anteriorly and laterally . . . .4. 

3. Anterior outer teeth disproportionately longer than the adjacent lateral teeth, 

slender, unicuspid and procurrent. . . Paralabidochromis gen. nov., p. 327 

Anterior outer teeth forming a graded series with the lateral teeth, not forwardly 

directed . . . . . . . . . . . Haplochromis 

4. Outer teeth enlarged and stout, with obliquely truncated and inwardly directed 

crowns ......... Macropleurodus, p. 299 

Outer teeth of both jaws small, bi- or unicuspid ; inner teeth in broad bands anteri- 
orly and laterally ........... 5. 

5. Lower jaw broad and flat, tooth bands in both jaws of uniform breadth antero- 

posteriorly ; those of lower jaw continued posteriorly on to the ascending part 

of the dentary ......... Hoplotilapia, p. 319 

Lower jaw stout, rounded anteriorly ; teeth aggregated anteriorly into two pyriform 

bands, contiguous at the symphysis ..... Platytaeniodus, p. 312 

Genus MACROPLEURODUS Regan, 1922 

Bayonia Boulenger, 191 1 {nee. Bocage 1865), type species Bayonia xenodonta Blgr. 
Macropleurodus Regan, 1922, Proc. zool. Soc, Lond. 189 ; type species : Haplochromis bicolor 

Boulenger, 1906, Ann. Mag. nat. Hist. (7) 17, 444. 
Haplochromis (part) ""] 

Paratilapia (part) > For references see synonymy under species. 

Hemitilapia (part) J 

Generic characters and synonymy 

Prior to Regan's revision of the Lake Victoria Cichlidae (Regan, 1922) specimens 
of the genus here recognized as Macropleurodus had been described as belonging to 
several genera and species. When Regan [op. cit.) defined the apparently new genus, 
Macropleurodus he did not have at his disposal specimens covering the wide range 
necessary to determine ontogenetic changes in certain characters. Consequently he 
failed to recognize that his genus was identical with Boulenger's Bayonia 
(Boulenger, 1911), and incorrectly assigned the type specimens of this and one other 
synonymous "species" to Haplochromis. 

Regan's diagnosis of Macropleurodus gave particular emphasis to the posterior 
premaxillary teeth : 

. . . several inner series of small teeth anteriorly and three or four series of 
enlarged teeth laterally, which are exposed when the mouth is shut" (Regan, 1922). 

ZOOL. 3, 7. 22§ 


The two specimens on which this description was based have the posterolateral 
inner teeth not only enlarged, but also similar in form to adjacent outer teeth. It 
is now known, however, that in small individuals these inner teeth are only slightly 
enlarged, if at all, and do not closely resemble the outer teeth. Specimens available 
indicate that most fish between 80 and 90 mm. standard length have a dentition 
intermediate between juvenile and adult types. In these, many of the typically 
juvenile, stoutly bi- and unicuspid inner teeth are replaced laterally by teeth 
differing only slightly in size and form from those of the outer series. Individuals 
below 80 mm. lack enlarged inner teeth. 

Thus, although Regan's diagnosis is incisive for fishes over 90 mm. S.L., it is not 
sufficiently comprehensive to include smaller individuals. 

On the other hand, the present collection shows that the morphology of the outer 
teeth is equally diagnostic and, moreover, is little affected by the size of the individual. 
Basically, the outer series is composed of stout teeth, having enlarged and obliquely 
truncated anterior cusps and disproportionately smaller posterior cusps. In the 
upper jaw these teeth are implanted obliquely to the long axis of the premaxilla, 
so that their crowns lie at an angle to it and the anterior cusp points inwards. Teeth 
of this type are present in all the specimens examined and are not known to occur 
in any Haplochromis or related species. 

The peculiar form of the outer teeth in Macropleurodus at once suggests affinity 
with Bayonia xenodonta Boulenger (191 1), in which the outer teeth are described as 
having " ... very large compressed crowns, with long anterior cusp directed 
inwards and very short or indistinct posterior cusp ... ".In this respect, the 
published figure of B. xenodonta (Boulenger, 191 1 and 1914) is somewhat misleading 
since the teeth are stouter than depicted. 

Through the courtesy of Dr. D. Guiglia, re-examination of the type specimen of 
B. xenodonta has been possible and has confirmed that the outer teeth are alike in 
Bayonia and Macropleurodus. Further, as Regan and Trewavas (1928) first observed, 
Boulenger's description of the inner teeth is inaccurate. Posteriorly these teeth 
form a double series, with six teeth on either side nearly as large as the outer, and 
in this respect conform to the dental pattern of young Macropleurodus. Also, the 
more anterior teeth of the two inner series are not " minute and conical " but are 
in fact tricuspid. Thickening of the buccal mucosa (probably a fixation artefact) 
has buried the inner teeth so that only their major cusps protrude. Two types of 
tricuspid teeth are present : one, the usual small tricuspid tooth found in many 
species of Haplochromis ; the other, stout and with the minor cusps displaced so as 
to form a triangular crown having the major cusp at its apex. Similar trigonid 
teeth are also found in Macropleurodus, both young and adult, but have not been 
observed in Haplochromis. 

There is also agreement in morphometric characters and, although Bayonia 
xenodonta has only twenty-four outer teeth in the upper jaw, this number is within 
the known lower range for Macropleurodus (see below). 

There seems little doubt, therefore, that in all salient features the type and unique 
specimen of Bayonia xenodonta is identical with small specimens of Macropleurodus 
bicolor, and that the two are conspecific. 


One other name has been given to cichlid fishes from Lake Victoria with teeth of 
the Macropleurodus type, namely Hemitilapia mater famili as Pellegrin, 191 3. I have 
examined the holotype (no. 12.278 in the collection of the Museum d'Histoire 
naturelle, Paris) and found it to agree closely with Macropleurodus of similar size. 
The teeth and dental pattern of the type and three other specimens represent an 
advanced stage in the transition from juvenile to adult condition ; that is, the 
postero-lateral premaxillary teeth of the first inner series are nearly comparable in 
size and form with the adjacent outer teeth. The remaining inner teeth are small 
and unequally bicuspid. 

Regan (1922) tentatively synonymized H. materfamilias with Haplochromis 
obliquidens Hilgendorf ; apparently he did not examine the type of H. materfamilias 
{op. cit., pp. 157 and 158) and was misled by Pellegrin's description. Comparison 
of the holotype with specimens of H. obliquidens at once reveals the existence of 
fundamental differences in the dentition of the two species. In H. obliquidens the 
movably implanted outer teeth are fine and numerous (50-70 in upper jaw) ; the 
posterior cusp is wanting, except very occasionally in small fish, whilst the anterior 
cusp, although obliquely truncate, is compressed and slender. In contradistinction, 
the immovable outer teeth of H. materfamilias are coarse and less numerous (24-40) ; 
with few exceptions a posterior cusp is present and the stout anterior cusp is 
circular in cross section. 


Cichlid fishes of the Haplochromis group as defined by Regan (1920, 1922) but 
differing from Haplochromis in having stout outer teeth with inwardly directed and 
obliquely truncated crowns ; anterior cusp long, slightly decurved and not com- 
pressed, the posterior cusp small and indistinct. Fishes over 90 mm. S.L. have one 
or more inner premaxillary tooth-series composed laterally of enlarged teeth similar 
in form to the adjacent outer teeth. Consequent upon the enlargement of the lateral 
teeth, the dentigerous surface of the premaxilla is broader laterally than anteriorly. 
In small individuals, where the inner teeth are small and bi- or unicuspid throughout 
the series, the outer teeth are already characteristic. Teeth usually exposed laterally, 
even when the mouth is shut. 

Macropleurodus bicolor (Boulenger) 1906 
(Fig. 1) 

Haplochromis bicolor (part) Blgr. 1906, Ann. Mag. nat. Hist. (7) 17, 444 (type only). 
Paratilapia bicolor (part) Blgr. 1907, Fish. Nile 479, pi. lxxxix, fig. 1 ; Idem, 191 1, Ann. Mus. 

Genova (3) 5, 68 ; Idem, 1915, Cat. Afr. Fish. 3, 346, fig. 234. 
Bayonia xenodonta Blgr. 191 1, Ann. Mus. Genova (3) 5, 70 ; Idem, 1915, op. cit., 488, fig. 338. 
Hemitilapia materfamilias Pellegrin, 1913, Bull. Soc. zool. France, 37, 313 ; Boulenger, 1915, 

op. cit., 492. 
Haplochromis obliquidens (part), Regan 1922, Proc. zool. Soc, London, 188. 
Macropleurodus bicolor, Regan, 1922, op. cit., 189. 


Fig. i.—Macropleurodus bicolor, holotype. 7/8 N.S. (From Boulenger, Fishes of the Nile. 

• • 


Cheek (?Head) 



• « 





• • • - 



• • 




• • 


• • • 






• • 




• • • • • 
••• . ••• 









. • • • ~**t 


• • 





• .• 






. ••• • 
• • •• • : "• • 









• • •• • • • 

• • • • 





• • • 



• • 


• •• * 






1 1 


— 1 

— , - ■ ■ 

50 60 70 80 90 100 110 1 

90 100 110 

Standard Length (mm.) 












• ••■■•.. - •.v^^vn..- 

Head (| SI) '• # * 

• • •» 

• ♦ • ♦ •••«•• a • 

LowerJaw(gHead) •• • • , 

25 : Eye (j Head) . ••./ ?. .• 

• •• .• 

• • ** • 


50 60 70 80 90 100 tfo 120 130 140 

Standard Length (mm.) 

Fig. 2. — Scatter-diagram to show individual variation and allometry of the characters 
indicated. The isometric relationship of head length and standard length is usual, as is 
the allometry of head length and the inter-related snout, eye and preorbital measurements . 
The negative allometry between length of lower jaw and headlength isunusual and seems 
to be related to the stronger jaws and enlarged teeth of larger specimens. These diagrams 
show scatter but not frequency. Within the size-range 100 to 125 mm. S.L., each dot in 
the denser aggregates represents at least two specimens. 


5, a). Posteriorly, the neurocranium does not differ significantly from the generalized 
type, except that the pharyngeal apophysis is not greatly depressed below the 
parasphenoid, and its pro-otic buttress is broad and bullate. 


Fig. 4. — Macropleurodus bicolor. (a) Neurocranium in left lateral view ; (b) dentary, 
(c) premaxilla, both in right lateral view. Skeleton prepared from a specimen of 
115 mm. S.L. 



Fig. 5. — Haplochromis michaeli. (a) Neurocranium, (b) dentary, left lateral view ; (c) 
dentary, occlusal view ; to show the skull and jaws of a generalized Haplochromis from 
Lake Victoria. Skeleton prepared from a specimen of 115 mm. S.L. 

The premaxilla shows considerable departure from the basic type found in most 
Haplochromis species. Whereas in Haplochromis there is slight ventral arching of 
the horizontal limb of this bone, in Macropleurodus the arch is greatly exaggerated, 
with its point of maximum curvature at the broadest part of the tooth band (Fig. 
4, b). Individuals with a marked dextrally developed tooth pattern have a cor- 
responding degree of asymmetry in the two halves of the premaxilla, which is then 
more acutely arched on the right side. 

ZOOL. 3, 7. 22§§ 


A comparative study of jaw musculature was made with Haplochromis sauvagei 
and H. prodromus, species in which the jaws show some deviation from the generalized 
Haplochromis condition towards that of Macropleurodus. Negligible differences 
were observed in the musculature of the three species, except that in M. bicolor the 
adductor mandibulae I measures only 28% of the head length, whilst in H. sauvagei 
and H. prodromus this muscle is 33% of head length. Adductor mandibulae II is also 
relatively shorter in M. bicolor (21-0% compared with 33-6% in H. sauvagei and 
26-9% in H. prodromus). In all other respects the cranial musculature of Macro- 
pleurodus is typical for a generalized Haplochromis species. 

Functional significance of the skull form, together with that of the well developed 
teeth, jaws and muscles, is best considered in relation to the predominantly molluscan 
diet of M. bicolor, and the manner in which its prey is secured. Whereas in most 
Haplochromis species the lower jaw is somewhat obliquely inclined when the mouth 
is shut, in M. bicolor it is almost horizontal. The position of the lower jaw at rest 
directly affects the manner in which the whole mouth is protruded ; in typical 
Haplochromis, maximal protrusion is achieved as a result of the mandible moving 
from an inclined to a horizontal position. On the other hand, when the mouth of 
M. bicolor is opened, mandibular movement is from the horizontal to a point below 
it. Premaxillary movement is also directed downwards by the near vertical ethmoid 
complex over which the premaxilla slides. In consequence, the whole mouth is 
protruded ventrally, with the upper jaw slightly in advance of the lower. The short 
and broad adductor mandibulae muscles allow the mouth to be retracted with 
remarkable rapidity, thereafter mounting a powerful and sustained pressure on any 
object held between the teeth. The crushing power of the jaws is further enhanced 
by the stout outer teeth and well developed bands of lateral and postero-lateral inner 

Aquarium observations show ^hat M. bicolor, when feeding, usually approaches a 
snail from above, rapidly protruding the mouth in an attempt to snatch its prey 
from the substrate. Once the snail is firmly held and suitably orientated — generally 
with the foot directly orally — there follows a series of short biting movements which 
crush the shell and thus free the soft parts, which alone are ingested. 

Attention has already been drawn to the similar skull structure in M. bicolor and 
H. prodromus ; it is not surprising therefore to find that both species have similar 
feeding habits. 

Ontogeny. Alizarin preparations of larval M. bicolor, H. macrops (a generalized 
species), and H. prodromus have been compared. These specimens reveal no funda- 
mental differences in osteology or dentition of the three species when compared at 
morphologically equivalent developmental stages. For example, at the latest stage 
examined (9-0 mm. total length ; yolk sac almost completely resorbed) the small 
conical outer teeth are morphologically and numerically identical in all three species. 
From these admittedly few observations it would seem that characteristic adult 
skull form and outer teeth must develop during post-larval ontogeny. 

Coloration and polychromatism 

Coloration in life. Adult females with greenish-yellow ground colour, becoming 


lighter or silver ventrally. Dorsal and anal fins yellow-green, the former with or 
without two irregular, dark, longitudinal stripes, the latter with two to four ill- 
defined yellow spots on the posterior part. Caudal and ventral fins generally colour- 
less, though the latter may sometimes be slightly dusky. Young of both sexes and 
sexually inactive males have similar coloration. 

Adult males (breeding coloration). Dark slate-blue ground colour, lighter, some- 
times silver, ventrally. Chest, branchiostegal membrane, lower jaw and ventral 
aspects of the cheek, black. Operculum and flank sometimes with a faint scarlet 
flush. Dorsal fin dusky, with deep red spots and streaks between the rays, especially 
intense on the soft part. Anal dusky, with well marked scarlet ocelli. Ventral fins 
black. Intensity of male coloration is correlated with sexual state ; inter-grades 
are known between the coloration described above and that of typical female or 
juvenile coloration. 

Besides normal sexually dimorphic coloration, certain fish exhibit a third colour 
pattern, in the form of an individually variable piebald, black on a yellow-green 
ground. The holotype is such a specimen (Fig. i). The bicolor pattern is clearly 
composed of vertically arranged irregular and often interrupted dark bands, which 
are generally continued across the body on to the vertical and paired fins. Although 
some are more intensely blotched than others, no intergrades are known between 
normal female coloration and bicolor variants. 

With two exceptions all bicolor individuals examined were females. The colour- 
pattern and degree of pigmentation differ in the two exceptional male fishes. Since 
protandry might be suspected, the gonads were sectioned and examined micro- 
scopically. In both fishes, however, there was evidence only of testicular tissue. In 
one fish the pattern is typical ; in the other the pattern is less intense and occurs 
on a darker ground than is typical for female fishes. 

Accurate frequency-estimates for bicolor individuals are difficult to obtain, since 
collectors show marked sampling bias in favour of these strikingly coloured fishes. 
However, in more rigorously controlled collections from one area, bicolor frequency 
amongst female fish in the 105 mm. to 125 mm. size class is approximately 30%, 
an incidence sufficiently high to justify regarding the phenomenon as being due to 
polymorphism and not to the maintenance of an atypical phenotype by recurrent 

If the two female colour patterns are accepted as an example of polymorphism, it 
is necessary, ex hypothesi, to consider the selective balance which must exist between 
the two forms. This question is further complicated by the apparently almost 
completely sex-limited polymorphism in M. bicolor. 

Since the genetical basis of polymorphism and sex determination is unknown 
for Macropleurodus, some hypothesis at least is desirable before considering the 
question of selective values for the two colour patterns. 

If, as in many fishes, the female is the heterogametic sex, then a possible (and 
doubtless oversimplified) explanation for this sex-limited polymorphism is that the 
gene or gene complex underlying development of a bicolor pattern may lie in a sex 
chromosome, be recessive to the gene or genes for normal colour, and be linked with 
a recessive lethal gene. Thus full expression of bicolor pattern could only be manifest 


in the heterogametic sex. Males carrying the double complement of recessive bicolor 
genes necessary for phenotypic expression in that sex, would, on the linkage supposi- 
tion, die as a result of simultaneously receiving the two recessive lethal genes. Since 
linkage is sometimes broken, a small percentage of male bicolor individuals might 
well be expected and the two bicolor male fishes in this collection are possibly such 

By this reasoning, either selection in favour of polymorph genes must be sufficient 
to compensate for loss of males and consequent unbalance of the sex-ratio or, 
alternatively, the unbalanced sex ratio may be the factor preventing spread of bicolor 
genes throughout the species, should these have a selective value slightly higher 
than " normal ". 

Two possible advantages associated with bicolor patterns, or genotypes, present 
themselves. Firstly, a bicolor pattern is, in effect, a disruptive one and may thus 
provide some protection against the attacks of predators. Studies on fish-eating 
birds (Cott, 1952) and on piscivorous fishes such as Bagrus and Clarias (personal 
observation) neither support nor negate this possibility, since M. bicolor has not 
been found among the prey of these animals. Secondly, there is the possibility, also 
unproven, that a female bicolor genotype, or a male heterozygous for bicolor, may 
possess some physiological advantage over other genotypes. 

Regrettably, then, insufficient positive evidence is available at present to warrant 
further discussion on the evolutionary aspects of polymorphism in Macropleurodus. 


Habitat. M. bicolor is widely distributed within Lake Victoria, occurring most 
frequently in littoral and sublittoral regions, especially where the bottom is hard 
(sand, rock or shingle), but only rarely over mud. Depth distribution is fairly 
restricted, with a maximum of between 30 and 40 feet (see also Graham, 1929). 

Food. From gut analyses of numerous specimens (throughout the size range 
60-150 mm.) it is apparent that snails and insect larvae are the predominant food 
organisms (see also Graham, 1929). Shell fragments are rarely found in the stomach 
or intestine although opercula are usually present. Aquarium observations confirm 
that almost the entire body of the snail is removed from its shell before ingestion 
takes place (vide p. 325), although small snails and, on occasion, thin-shelled species, 
may be crushed intra-orally before being swallowed. As a result of this feeding 
mechanism snail remains are so fragmentary as to preclude accurate identification ; 
remains of Gabbia sp. have, however, been positively identified on several occasions. 

The insects most commonly recorded from the pabulum of M. bicolor are larvae 
of the boring may-fly, Povilla adusta Navas, with other larval Ephemeroptera, and 
larval Chironomidae occurring less frequently. The proportion of insect to molluscan 
food eaten is difficult to determine and is probably related to local and cyclic abund- 
ance of these organisms. 

In the light of numerous gut analyses which are now available for this species, 
Graham's record (1929) of fish and cichlid eggs from the stomach of M. bicolor requires 
some comment. Apart from this record no other instances of piscivorous habits are 
known for M. bicolor. Unfortunately, Graham does not give a detailed analysis of 


the gut contents, particularly of the number of specimens from which his data are 
derived. Bearing in mind this limitation, it is suggested that the fish-remains and 
eggs were from the stomachs of different individuals and consequent upon brooding 
female fishes swallowing their own young, a relatively common occurrence when the 
fishes are caught in gill-nets. 

Breeding. The exact spawning grounds of M. bicolor are unknown : eggs and 
larvae at all stages of development have been found in the mouths of various female 
fishes and it is presumed that the species is a mouth-brooder. Brooding females 
have been caught in all parts of the species range. The smallest sexually active fish 
examined was a female 96 mm. long [ex Kisumu). The habitat of post-larval fishes 
is unknown. 


Unlike the other monotypic genera M. bicolor can apparently be related to an 
extant species of Haplochromis. Similarity in skull architecture of M. bicolor and 
H. prodromus has already been noted. There are additional similarities in the short 
and stoutly constructed jaws of both species, besides a marked resemblance in 
general facies. Against these resemblances must be set the very different tooth form 
and dental pattern of H. prodromus, although in this species the teeth are stout and 
the inner series well developed. The structure of the head and dental patterns 
suggest that, at a functional level, the condition represented by H. prodromus might 
well be considered pre-adaptive to the development of a relatively massive dentition, 
such as that of M. bicolor. 

Study material and distribution records 

Museum and Reg. No. 

S.L. (mm.). 



British Museum (N.H.) : 

1906. 5. 30. 414 (type of H. bicolor) 

1906.5.30.378 .... 


1928.5.24. 1-3 .... 



Bunjako (Uganda) 


See below 



. M. Graham 

Paris Museum : 

12.278 (holotype of H. mater- 
familias ..... 

12.279-281 (paratypes of H. mater- 
familias ; proportions not in- 
cluded in description above) 


Port Florence, 


. Alluaud and 


Genoa Museum : 

(Type of Bayonia xenodonta Blgr.) . 




B.M.(N.H.) : 

1955.2. 10. 5-11 .... 



. E.A.F.R.O. 

1955.2. 10.50-57 


Jinja, Napoleon 


1955.2. 10.43-46 


. Beach nr. Nasu Point, 
Buvuma Channel 



Museum and Reg. No. 

S.L. (mm.). 



B.M.(N.H.) :— cont. 

1955.2. 10.22-23 

82 and 97 

Grant Bay (Uganda) . 


1955.2. 10.21 


Dagusi Island 


1955.2. 10.65-73 


. Off southern tip of 
Buvuma Is. 


1955.2. 10.24-42 and 74-83 . 


. Harbour at Entebbe . 


1955.2. 10.58-62 


Bugonga Beach 


1955.2. 10.12, 16-20 and 48 . 


Busongwe Bay 
(Kagera River mouth) 


1955.2. 10.63-64 

. no and 147 

Ma jit a Beach 
(Tanganyika Terr.) 

i > 

1955.2. 10.4, 49 and 84 




Graham (1929) records the occurrence of M. bicolor as follows : 

Kenya : Kavirondo Gulf : Off Sukuri Island. 

Off Ulambwi bay. 
Mbita Passage. 
Near Nzoia River. 
Kadimu Bay. 
Tanganyika Territory : Mwanza. 

Smith Sound. 

Genus PLATYTAENIODUS Boulenger, 1906 

Platytaeniodus Boulenger, 1906, Ann. Mag. nat. Hist. (7) 17, 451 ; Idem, 1907, Fish. Nile, 493 ; 

Idem, 1915, Cat. Afr. Fish. 3, 426, fig. 292 ; Regan, 1922, Proc. zool. Soc, Lond. 190. Type 

species : Platytaeniodus degeni Blgr. 1906. 

Generic synonyms : 

Astatotilapia (part) "\ ^ , , 

tt u 7 • / 1\ >For references see synonymy under species. 

Haplochromis (part) J J J J r 

Generic characters and synonomy 

Both Boulenger and Regan considered the premaxilla and its tooth pattern 
diagnostic. Boulenger (1914) states : 

"... the alveolar surface of the premaxilla widening towards the pharynx, the 
band of teeth in the upper jaw horseshoe shaped ..." 

Additional material shows, however that in fishes below 100 mm. standard length 
the posterior premaxillary dentigerous (alveolar) surfaces are not always expanded 
medially. Nevertheless, even in small specimens the premaxilla is stouter and its 
dentigerous surface wider than in Haplochromis ; posteriorly the teeth are arranged 
in several rows, so that the premaxillary tooth band is always clearly U-shaped, 
with the arms at least as broad as the medial part. Broadening of the posterior 
alveolar surfaces is gradual and shows positive allometry with standard length ; in 
some large individuals the left and right surfaces are closely apposed in the mid-line 
(Fig. 6, c). 


Tooth bands in the lower jaw are more readily diagnostic and less subject to 
variation with absolute size than those of the premaxilla. In Platytaeniodus the 
mandibular teeth are confined to the anterior and antero-lateral portions of the 
dentary and are grouped into two broad and roughly pyriform patches, contiguous 
at the symphysis ; posteriorly there is a short, single row of four to seven teeth 



Fig. 6. — Platytaeniodus degeni. (a) Premaxilla, (b) dentary, both in occlusal view. 
Skeleton prepared from a specimen 80 mm. S.L. (c) Mouth of the holotype, ca. x 4. 
[(c) from Boulenger, Fishes of the Nile.'] 


lying between the ascending part of the ramus and the anterior tooth bands (Fig. 6, 
b and c). 

When provisionally referring Astatotilapia jeanneli Pellegrin to Haplochromis 
macrops Blgr., Regan (1922) was apparently misled by the large eye and shallow 
preorbital of A. jeanneli. There is undoubtedly some resemblance between H. 
macrops and P. degeni, but this is confined to superficial characters, and is belied 
by their fundamentally different dentition. Re-examination of the types of A. 
jeanneli reveals that the premaxillary teeth are arranged in four or five series of equal 
breadth both laterally and posteriorly, a condition never observed in H. macrops. 
Furthermore A . jeanneli has the mandibular teeth grouped anteriorly and laterally 
in five series, with only a short, single series posteriorly. That is to say, both type 
specimens have a dentition typical for small P. degeni. In other characters too, 
notably the mouth with its broad lower j aw, shorter than the upper, and the almost 
completely hidden maxilla, A . jeanneli agrees more closely with P. degeni than does 
H. macrops or any other Haplochromis species. 

On these grounds, therefore, I consider A. jeanneli to be synonomous with P. 


Cichlid fishes of the Haplochromis group, but differing from Haplochromis in 
having broad bands of teeth on the posterior part of the premaxillary dentigerous 
surface, which is expanded medially in large specimens but is of almost equal breadth 
anteriorly and laterally in fishes of less than 100 mm. S.L. Teeth on the dentary 
grouped into two, broad, pyriform, curved and contiguous bands anteriorly and 
antero-laterally, but continued posteriorly as a short single row only. Lower jaw 
usually shorter than the upper ; maxilla almost completely hidden below the 

Fig. 7. — Platytaeniodus degeni, £. Drawn by Miss L. Buswell, 


Platytaeniodus degeni Boulenger, 1906 

(Fig. 7) 

Platytaeniodus degeni Boulenger, 1906, I.e.; 1907, I.e. and pi. xci, fig. 1; 1915, I.e. and fig. 292; 

Regan, 1922, 190, fig. 14. 
Astatotilapia jeanneli Pellegrin, 1913, Bull. Soc. zool. France, 37, 313. 
Haplochromis jeannelli (Pellegrin), Blgr. 1915, Cat. Afr. Fish. 3, 291. 
Haplochromis maerops (part) Regan, 1922, Proc. zool. Soc. London, 166. 


Thirty-six specimens (size range 67 to 154 mm.) comprising the type and other 
specimens in the British Museum (Nat. Hist.) (including material newly collected 
by E.A.F.R.O.), as well as the types of A. jeanneli are considered in this description. 

Since most characters tabulated below show some allometry with standard length 
these data are grouped into two size classes. In some characters intra-group vari- 
ability is high, but further subdivision into smaller groups is impracticable. 

In the table of proportions, head length, depth of body and length of caudal 
peduncle are expressed as percentages of standard length ; all other characters are 
expressed as percentages of head length. Range and mean are given for each 

Table I. 

Standard Length. 
Depth of body . 

Length of head 

Depth of preorbital . 

Least interorbital width 

Length of snout 

Diameter of eye 

Depth of cheek 

Length of lower jaw . 

Caudal peduncle 

67-93 mm. 
(23 specimens) 

M =35-6 
M = 32-4 
M = 14-2 
M = 27-1 
M = 30-0 

M = 30-8 
M = 19-7 
M = 37-0 
M = 16-5 

98-154 mm. 
(13 specimens). 

M = 38-4 
M = 32-7 
M = 15-2 
M = 31-2 

M = 34-8 
M = 24-8 
M = 23-4 
M = 36-0 
M = 16-7 

Dorsal profile of head and snout gently (in a single specimen somewhat strongly) 
decurved ; mouth horizontal ; lower jaw equal to or more usually shorter than upper, 
its length /breadth ratio i-i6-i-68 : in some specimens the lower jaw, including 
lips, is slightly broader than the upper. Lips well developed and somewhat thickened ; 
maxilla almost completely hidden beneath the preorbital, with only its postero- 
ventral tip exposed and extending to below the anterior orbital margin, or slightly 


The holotype, a male of 115-0 mm. S.L., is figured by Boulenger (1907 and 1915). 
This fish is both somewhat atypical and slightly distorted in preservation and the 
impression given of a deep ventral profile and a slightly oblique mouth is not 

Gill rakers. Short and stout, 7-8 on lower limb of the anterior arch, the lowest one 
or two usually reduced. 

Dentition. Premaxilla with 4-8 rows of teeth ; dentary with 4-6 rows. 

With few exceptions, specimens over 100 mm. S.L. have the posterior part of the 
premaxillary dentigerous surface expanded medially, so that the upper tooth bands 
are broader posteriorly than anteriorly. In small individuals, although there are 
several series of teeth posteriorly, the tooth band is either of equal width at all 
points, or medial expansion of the premaxilla may have begun, causing the band to 
be very slightly wider posteriorly (Fig. 6, a). Expansion of the posterior surface is 
not correlated with an increase in the number of tooth rows borne on it, which are, 
in fact, equal to or slightly fewer than those on the anterior part of the premaxilla. 

Mandibular dental pattern as described for the genus ; pyriform band from half 
to two thirds as broad as long. 

Teeth are variable in form, those of the outer series slenderly conical, with or 
without an admixture of unequally bicuspid teeth. Teeth of the inner series all 
unicuspid in most specimens above 100 mm. S.L. and in some below that size ; 
otherwise the outermost teeth unicuspid and the remainder tricuspid or exceptionally, 

Scales ctenoid ; lateral line interrupted, with 31 (f.2), 32 (f.5), 33 (f.14), 34 (, 
35 (f.2), or 36 (f.3) scales. Cheek with 3-4 series of imbricating scales ; 7-9 scales 
between lateral line and origin of dorsal fin ; 7-9 (rarely 10) between the pectoral 
and pelvic fins. 

Fins. Dorsal with 24 (f.3), 25 (f.23) or 26 ( rays, anal with 11 (f.6), 12 (f.23) 
or 13 (f.7) rays, comprising XV-XVII 8-10 and III 8-10 spinous and soft rays. 
Pectoral fin 22-6-31-8 (M = 28-0) per cent of standard length. Caudal truncate, 
scaled on its proximal half only. Pelvics with the first ray produced, extending to 
the vent in a few specimens and to the spinous part of the anal in most. 

Syncranium. Since the form of premaxilla and dentary in P. degeni is correlated 
with the well-developed tooth pattern, both these bones depart very strikingly from 
the typical Haplochromis condition. 

In small individuals of P. degeni, the premaxilla bears a superficial resemblance 
to that of Hoplotilapia, particularly with regard to the dental pattern, but in large 
fishes it is unique. On the other hand, the dental pattern and morphology of the 
dentary are comparable in both large and small individuals. The dentary is charac- 
terized by its broad and laterally expanded anterior tooth-bearing portion, which 
imparts to this bone an appearance unique amongst the Lake Victoria cichlids. 

The neurocranium of P. degeni is intermediate between the generalized Haplo- 
chromis type and that of M. bicolor. It is strictly comparable with the neurocrania 
of species of the H. crassilabris group. Here the skull is characterized by a somewhat 
shortened and steeply inclined ethmo-vomer complex ; in consequence, the anterior 
skull profile is also steep. In a typical Haplochromis skull the ethmo-vomer is longer 


and rises less steeply, meeting the downward sloping frontals at a wide angle. As a 
result, the anterior profile is shallower and also more acute than in the " crassilabris " 
type skull (Fig. 5). 

Jaw musculature in P. degeni dos not differ greatly from that of a generalized 
Haplochromis species. 

Coloration. Preserved material. Males : dusky to dark grey ; dorsal and anal 
fins dark ; pel vies black, caudal colourless. Females and immature individuals : 
silver-grey or light brown ; fins colourless. 

Transverse and longitudinal banding sometimes occurs, being most clearly marked 
in females and young individuals ; when present, there is a well marked median 
longitudinal stripe, a fainter and interrupted band running slightly below the dorsal 
fin, and eight or nine narrow transverse stripes on the flank and caudal peduncle. 
Faint lachrymal and interocular stripes may also be present. The presence and 
intensity of these markings is apparently related to the emotional state of the fish 
or may only appear after death. 

Coloration in life. Sexually active males : ground colour light blue-grey, lips 
iridescent blue. Chest and branchiostegal membrane black. Fins ; dorsal sooty, 
lappets and spots on soft part red ; caudal with red flush, most intense along margin ; 
anal with dusky pink flush and several yellow ocelli. Females : ground colour 
golden fawn ; all fins neutral, dorsal with orange lappets and spots, especially on 
the soft part ; caudal with orange margin and maculae ; anal with faint or well 
marked yellow ocelli. 


P. degeni is recorded from several areas in Lake Victoria (see below), but as so 
few specimens are known it is not possible to generalize on habitat preferences. 
From the scanty data available it appears that the species is probably restricted to 
littoral and sub-littoral regions where the water is less than fifty feet deep. Specimens 
have been caught in nets set over both hard and soft substrates, but the greater 
number came from stations having a ! sand or shingle bottom. 

Food. The distinctive dentition of this species suggests a highly specialized diet. 
Tantalizingly few fish, however, have yielded ingested material. Twelve specimens 
have been examined, all of which were caught in nets set overnight or in seines 
operated during varied daylight hours. According to the substrate over which they 
were living, ten fishes had either sand grains or organic mud in the stomach and 
intestines, together with fairly dense aggregations of mucus. Two fishes, caught on 
different occasions at a station near the southern tip of Ramafuta Island (Buvuma 
Channel), had the entire alimentary tract filled with the diatom Melosira. Diatoms 
from the stomachs of these fishes showed only slight signs of digestion, but samples 
taken from the mid-intestines and recta were almost completely digested. Animal 
remains, occurring sporadically, included insect larvae, Hydracarina, fragments of 
Copepoda, Ostracoda, and in two specimens shell fragments of Pelecypoda (Sphae- 

Most guts also contained some diatoms and blue-green algae, the former apparently 
digested, the latter intact. The very small quantity of ingested material in any one 


individual is striking and no particular organism, or group of organisms, occurs with 
sufficient frequency to indicate what the food of P. degeni may be. Since sand and 
bottom debris is significant in the majority of specimens, it is possible that the species 
may feed on the micro-fauna and flora living on and within the substrate. Thus, 
broad bands of jaw teeth may serve to rasp and loosen food from the surrounding 

Breeding. Spawning sites are unknown ; only two females, both from beaches in 
the Mwanza area, have been found with eggs in the mouth. The smallest individual 
with demonstrably active gonads was a female, 71-0 mm. long. 


There is no obvious relationship between P. degeni and any known Lake Victoria 
species or species-group of Haplochromis, with which genus the species shows 
fundamental affinities. The peculiar premaxillary and mandibular tooth patterns 
serve to set P. degeni apart from even those Haplochromis with several series of 
inner teeth. Regan (1922) considered P. degeni as being " very near " to H. prod- 
romus, which species he believed " shows a slight departure from the normal Haplo- 
chromis dentition towards the Platytaeniodus type ". His opinion was based on the 
holotype and then unique specimen of H. prodromus. Summarizing unpublished 
data on H. prodromus, it is clear that the type specimen has an aberrant dental 
pattern and that its resemblance to P. degeni is purely superficial. Whereas in large 
Platytaeniodus there is an actual expansion of tooth-bearing surfaces, in H. prodromus 
only the tooth-band is apparently expanded ; its increased breadth is actually due 
to the posterior teeth being more widely separated from one another than are the 
anterior teeth. In no specimen of H. prodromus is the upper tooth band as broad 
posteriorly as anteriorly, yet this is the usual condition in P. degeni. Further, the 
dentary of P. degeni differs considerably from that of H. prodromus. The evolution 
of a wholly multi-seriate dentition has probably occurred more than once within the 
Lake Victoria species-flock, as for instance in the H. sauvagei group and again in 
the monotypic genera. Thus any apparent relationship between P. degeni and H. 
prodromus should be considered as consequent upon convergent evolutionary trends, 
the ultimate expressions of which are achieved by manifestly dissimilar means. 

Study material and distribution records 

Museum and Reg. No. 

S.L. (mm.). 



British Museum (Nat. Hist.) : 

1906. 5. 30. 51 1 (holotype) 


. Bunjako (Uganda) 




Sesse Is. (Uganda) 

1928.5.24 .... 


Mbita Passage 

. M. Graham 

Paris Museum : 

12 . 262 (holotype of A . jeanneli) 


Port Florence 

. Alluaud and 



12 . 262 (paratype of A . jeanneli) 





Museum and Reg. No. S.L. (mm.). ' Locality. Collector. 

B.M. (N.H.) : 

1955.2. 10.91-94 .... 73-79 • Kisumu (Kenya) . E.A.F.R.O. 

1955-2.10.105-106 . . . 78-90 . Kamarenga (Kenya) . Ditto 

1955-2. 10. 115 .... 74 Kendu (Kenya) 

1955.2. 10.88 .... 74 Likungu (Kenya) . „ 

Genus HOPLOTILAPIA Hilgendorf, 1888 

Hoplotilapia Hilgendorf, 1888, S.B. Ges. naturf. Fr. Berlin, 76-77 (type species (Paratilapia ?) 

retrodens Hilgendorf, I.e.) ; Regan, 1922, Proc. zool. Soc, Lond. 190. 
Cnestrostoma Regan, 1920, Ann. Mag. nat. Hist. (9) 5, footnote p. 45 (type species Paratilapia 

polyodon, Blgr.). 
Haplochromis (part) "] 

Paratilapia (part) }■ For references see synonymy under species below. 
Hemichromis (part) J 

Generic characters and synonymy 

The holotype of Hoplotilapia retrodens (in the collections of the Zoologisches 
Museum der Humboldt-Universitat, Berlin) has not been examined by me, nor can it 
be definitely established whether this specimen is still in existence. Dr. Kurt Deckert 
of the Zoologisches Museum has, on two occasions, kindly attempted to locate 
several specimens, including the type of H. retrodens. Of these he writes : " Ich 
muss Ihnen leider mittheilen, dass unser Suchen nach den verlangten Typen ohne 
Erfolg geblieben ist, obwohl ich mit grosser Sicherheit anneheme, dass sie nicht 
verloren-gegangen sind. ' ' 

Hilgendorf s original description of {Paratilapia?) retrodens (1888), although brief 
and lacking detail, nevertheless stresses characters which clearly separate this 
species from others of the Haplochromis group : viz. a multi-seriate dental pattern 
with stout and enlarged posterior teeth. Pfeffer's redescription (1896) of the same 
specimen confirms and extends this account. In the material at my disposal, how- 
ever, the posterior teeth are clearly enlarged only in the upper jaw, and not in both, 
as stated for the type. Specimens described below agree closely with the type in 
the other characters described by Hilgendorf and Pfeffer ; slight differences in scale 
numbers can probably be attributed to different methods of making these counts. 

In addition to the four specimens of H. retrodens in the British Museum (Nat. 
Hist.) it has been possible, through the kindness of Dr. Delfa Guiglia, to examine 
the type of Paratilapia polyodon Blgr. and one other specimen (Museo Civico 
di Storia Naturale, Genoa, reg. no. G.E. 12.994) determined by Boulenger as 
P. polyodon, and to confirm Regan's view (1922) that these are conspecific with 
H. retrodens. 


Differing from Haplochromis as defined by Regan (1920 and 19226) in having 
broad bands of teeth in both jaws, well developed and usually of almost uniform 
breadth throughout or very slightly narrower posteriorly. Posterior teeth of the 
upper jaw enlarged and stout, those of the lower jaw slightly, if at all, enlarged, but 


the tooth-band continued posteriorly on to the steep ascending contour of the 
dentary. Lower jaw wide and flat, almost square in anterior outline, slightly shorter 
than the upper. 

Since only three of the collected specimens of this genus are small it is not possible 
to generalize on differences which apparently exist between the dentition of adult 


Fig. 8. — Hoplotilapia retrodens. (a) Neurocranium, (b) dentary in lateral view, 
(c) dentary, occlusal view. Dentition only part indicated. From a specimen of 125 
mm. S.L. 


and juvenile fishes. In these three specimens (74-0, 76-0 and 55-0 mm. S.L.) the 
tooth bands are broad anteriorly, being composed of 5, 3, and 3 series respectively. 
Laterally, however, they are reduced to two series, whilst postero-laterally only the 
outer series persists. In none is the dentition continued onto the ascending part of 
the dentary, although the shape of the lower jaw is as in the adult. 

Two adults of 96 and no mm. S.L., collected in a single seine haul at Bukakata, 
retain these presumedly juvenile dental characters. They form a graded morpho- 
logical series with a third specimen (134-0 mm. S.L., from the same station) which 
exhibits only slight departure from the " typical " condition. 

The broad and shallow lower jaw (Fig. 8, Band c) of typical individuals is unique 
amongst Lake Victoria Cichlidae. A few specimens of H. retrodens have the dentary, 
at least in external appearance, similar to that of Haplochromis, although in every 
case the dental pattern is typical for Hoplotilapia. 

Fig. 9. — Hoplotilapia retrodens, $. Drawn by Miss L. Buswell. 

Hoplotilapia retrodens Hilgendorf, 1888 
Fig. 9. 

(Paratilapia ?) retrodens Hilgendorf, 1888, S.B. Ges. naturf. Fr. Berlin, 76. 

Hoplotilapia retrodens Hilgendorf, t.c, 77. 

Hemichromis retrodens, Pfeffer, 1896, Thierw. O. Afr. Fische, 19. 

Haplochromis bicolor (part) Boulenger, 1906, Ann. Mag. nat. Hist. (7) 17, 444 ; two specimens 

B.M. no. 1906. 5. 30. 417 and 418. 
Paratilapia bicolor (part) Boulenger 1915, Cat. Afr. Fish. 3, 346. 
Paratilapia poly don Boulenger, 1909, Ann. Mus. Genova (3) 4, 306, rig. ; Idem, 191 1, ibid. 5, 

68 ; Idem 1915, Cat. Afr. Fish. 3, 349, fig. 236. 
Cnestrostoma polyodon (Boulenger), Regan, 1920, Ann. Mag. nat. Hist. (9) 5, footnote p. 45. 



From available material it would appear that only the interorbital width shows 
marked allometry with standard length. It must, however, be borne in mind that 
paucity of specimens within the smaller and larger size groups may obscure such 
relationships, especialy since, in those size-groups which are well represented, 
individual variability is high. With the exception therefore of interorbital width, 
ranges and means are given for the sample as a whole. For interorbital width the 
range and mean are given for three size groups, 74-115 mm. (N = 21), 116-130 mm. 
(N = 32), and 131-144 mm. (N = 11). 

Depth of body 31-6-41-6 (M = 38-3), length of head 30-1-34-8 (M = 32-0) per 
cent of standard length. Dorsal profile of head and snout straight or slightly concave, 
steeply sloping. Preorbital depth 12-5-18-8 (M = 16-4) per cent of head length ; 
least width of interorbital 24-2-33-8 (M = 28-7), 28-6-35-1 (M = 31-5) and 31-1- 
33-4 (M = 31*6) per cent for the three size groups respectively ; length of snout 
29-0-36-8 (M = 32-5), eye 23-8-29-6 (M = 26-9), depth of cheek 19-5-28-2 (M = 
23-9) per cent of head length. 

Lower jaw slightly shorter than upper 33-7-40-8 (M = 38-2) per cent of head, the 
length /breadth ratio from broader than long to 1-33 times as long as broad. Mouth 
horizontal, lips somewhat thickened. Posterior tip of the maxilla extending to the 
vertical from the anterior margin of the orbit or as far as the pupil. 

Described from 64 specimens, 55-144 mm. standard length. 

With few exceptions, there is remarkable uniformity in the general facies of H. 
retrodens. In this respect the figure of Paratilapia polyodon type specimen in 
Boulenger (1909 and 1915) can be considered fairly representative. The lower jaw 
in this specimen has, however, been broken and subsequently distorted in preserva- 
tion, consequently the ventral head profile of the figured specimen is inclined upward 
and is not horizontal, as it would be in life. The greatest departure from typical 
physiognomy and body form is seen in a single specimen from Bukoba (Tanganyika) 
which has the body relatively elongate and the head profile strongly curved. Despite 
this aberrance in gross morphology, the dentition and other fundamental characters 
of this specimen are typical. 

Teeth and dental pattern. Fishes above 90 mm. S.L. have 5 to 8 series of teeth 
anteriorly and 4 to 5 series posteriorly in the upper jaw ; 5 to 8 (rarely 9 or 10) 
series anteriorly and 3 to 5 series posteriorly in the lower jaw. From 40 to 68 teeth 
in outer series of the upper jaw. Teeth small, those of the outermost series largest, 
variable in form, but usually unicuspid ; some bi- and tricuspid teeth occur in the 
inner series. Two specimens from Majita and Mwanza (Tanganyika Territory) have 
markedly aberrant, stoutly conical or nearly molariform teeth in all series of both 
jaws. In both specimens the dental pattern is otherwise typical. 

Syncranium. The premaxilla is comparable in form with that of Haplochromis, 
except that it is stouter, especially posteriorly, and its dentigerous surface broader. 

As noted above, the broad and greatly flattened dentary is unique amongst Lake 
Victoria cichlids (Fig. 8). 

The neurocranium agrees closely with that of Platytaeniodus and with Haplo- 
chromis of the " crassilabris " species-group (see p. 316). 


Pharyngeal bone with a triangular and approximately equilateral dentigerous 
surface ; pharyngeal teeth cuspidate, the median series often enlarged. 

Scales ctenoid, lateral line interrupted, with 31 (f.i), 32 (, 33 (f.38), 34 (f.14) 
or 35 (f.i) scales. (Hilgendorf (1888) gives L.S. 30 [probably on a mid-lateral series] 
for the holotype.) Cheek with three or four series of imbricating scales. (Hilgendorf 
[loc. cit.) gives 4-5 rows) ; y-S scales — rarely 6 — between lateral line and origin of 
dorsal fin ; 8-9, less commonly 7, between pectoral and pelvic fin bases. 

Fins. Dorsal with 24 (f.4), 25 (f.45) or 26 (f.13) rays, comprising XV-XVII 8-10 
spinous and soft rays. Two specimens have the formulae XIV 8 and XVII 6, but 
there are indications that these are the results of wounding and subsequent irregular 
healing. Anal with n (f.6), 12 (f.54) or 13 (f.4) rays, i.e. Ill, 8-10. Caudal fin trun- 
cate, scaled on proximal half only. Pectoral fin 23-9-33-0 (M = 28-4) per cent of 
standard length. Pelvics with first ray produced in both sexes but proportionately 
longer in adult males. 

Coloration. Colours of preserved specimens are variable. Adult males : dark 
grey to black, median fins dark, caudal colourless or maculate ; pelvics black. 
Females and immature males : light olive brown to silver, all fins yellow or colour- 
less. Sometimes faint traces of transverse bars, an interrupted band below insertion 
of dorsal fin and a well-marked mid-lateral stripe. 

Colours in life. Adult males : ground colour dark grey-green (darkening to deep 
slate-grey immediately after death), chest and branchiostegal membrane black. 
Dorsal fin dark grey-green, lappets red ; red spots, often coalescing into streaks, 
between the rays. Caudal with proximal third to half black, distal part red. Anal 
with dark base, remainder red ; ocelli yellow. Pectoral colourless ; pelvics black, 
with faint red flush along median rays. Adult females : ground colour light olivaceous 
dorsally, shading through sulphur-yellow to pearly-white ventrally ; usually two 
longitudinal bands as described above, of variable intensity. Lips yellow. Dorsal 
and caudal fins olivaceous ; anal colourless or with slight yellow flush ; small and 
ill-defined orange spots sometimes occur in the position of the male ocelli. Pelvics 
and pectoral fins yellow. 

Three specimens (2 ?, 1 <J) from Bukakata, which were noted when discussing 
atypical dental patterns (p. 321), also exhibit aberrant coloration. This is known 
only from preserved material, in which it has the form of three large and elongate 
black spots, arranged mid-laterally, on a light ground colour. 

Almost completely sex-limited polychromatism occurs in this species and, as in 
M. bicolor, it is the female which usually exhibits atypical coloration. Besides the 
black and yellow piebald, as described for M. bicolor, there is a second and more 
colourful, if less distinctive, pattern. Any attempt to describe this pattern must 
perforce be imprecise, since intensity and detail show a remarkable range of indi- 
vidual variability. The ground colour is invariably a light sandy-yellow, with a 
superimposed orange flush, usually most intense on the head. Dorsally there are a 
number of irregular and ill-defined dark blotches, separate or confluent, which occa- 
sionally extend on to the flank. In some individuals this dorsal pigmentation is 
comparable with the clearly defined blotches of a typical bicolor pattern ; in others 
it is more diffuse and individual elements are only faintly discernible. The whole 


body, including the fins, is also peppered with small melanophores, particularly 
on the head and opercula. Fin coloration is extremely variable, but the caudal and 
anal are usually flushed with bright orange. 

Sampling bias in favour of atypically-coloured individuals undoubtedly occurs 
and precludes the accurate estimation of frequency. Of thirty-five females examined, 
thirteen were " bicolor " and eleven of the other pattern. Polychromatism is known 
in populations from most areas in which the species has been collected. 

A single male (119-0 mm. S.L., Busongwe, Kagera River area) had an incipient 
bicolor pattern, resembling in appearance and degree of pigmentation one of the 
" bicolor " males of M. bicolor (p. 309). 

The genetical basis of, and selection factors maintaining polychromatism in H. 
retrodens remain undetermined, as previously explained in the case of M . bicolor. 


H. retrodens is widespread throughout Lake Victoria (see below). Sampling in 
many habitats shows the species to be restricted to littoral and sub-littoral areas, 
especially where the substrate is hard, and usually where there is submerged vegeta- 
tion. The majority of specimens was caught in water from twelve to twenty feet 
deep, with a few from slightly deeper water. 

Food. The gut contents of sixty-five individuals (size-range 75-0-144-0 mm. S.L.) 
have been examined ; of these, only seven were empty. Mollusca (particularly 
Lamellibranchiata) are the predominating food, being recorded from thirty-six 
fishes. The majority of Mollusca is represented by finety broken shells. Insecta 
(f.8) — especially Ephemeroptera, Trichoptera and Chironomidae — together with 
Crustacea (f.3) and Hydracarina (f.2) occur less frequently. In fifteen fishes the 
ingested material consisted almost entirely of fragmented plant epidermis, whilst 
eleven others contained only sand grains or organic mud. 

Observations made on fishes living in aquaria show that H. retrodens, when feeding, 
repeatedly makes short, darting movements into or over the substrate, the broad 
and horizontal mouth serving as a scoop or shovel. Much bottom material is spilled 
from the mouth or with the exhalent current. Snails were retained and apparently 
crushed within the jaws before being passed back to the pharynx. The presence of 
macerated plant epidermis in several fishes may be explained either as ingested 
bottom debris, or as the result of the fish actively scraping the leaves and stems of 
submerged plants, for which purpose the multi-seriate dentition would seem adapted. 
It may be significant to note that the frustules of epiphytic diatoms found in the 
stomachs of these fishes were empty, as were the epidermal cells which had been 

Breeding. No information is available on the breeding habits of H. retrodens. The 
smallest sexually mature individual was a female of 96 mm. S.L. (Bukakata, Uganda). 


Morphologically, Hoplotilapia retrodens, like P. degeni is relatively far removed 
from any Lake Victoria Haplochromis species or species group. Neither is it closely 


related to the other monotypic genera of that lake. Regan's remark (1922, p. 159) 
that " A remarkable group of three species includes Haplochromis sauvagei and the 
monotypic genera Macropleurodus and Hoplotilapia, which scarcely differ from each 
other except for the considerable differences in dentition, ... "is difficult to 
endorse, unless the various structures whose forms are modified in association with 
the dentition and feeding habits are included in the term " dentition ". Macro- 
pleurodus and Hoplotilapia have manifestly dissimilar jaw morphology and also 
differ in the form of the neurocranium. These structural differences impart a charac- 
teristic physiognomy to the species, which allows them to be recognized without 
reference to the dental form and pattern. The occurrence of " bicolor " patterns in 
both M. bicolor and H. retrodens is suggestive, but at present little importance can 
be attached to this character since it occurs in at least three widely divergent species 
of Haplochromis as well as in M. bicolor. If neurocranial form can be considered as 
being of phylogenetic value, then the affinities of Hoplotilapia lie with the H. crassi- 
labris species group, although H. retrodens has departed considerably from this 
complex in the form of its jaws and dental pattern. 

Hoplotilapia and Macropleurodus show interesting trophic parallels, both between 
the two genera and in relation to the three mollusc-eating cichlids of Lake Victoria, 
viz. Astatoreochromis alluaudi, Haplochromis pharyngomylus and H. ishmaeli. 
Ecologically, Hoplotilapia and Macropleurodus occur together with A. alluaudi and 
H. pharyngomylus, but the diet of Hoplotilapia and especially of Macropleurodus, 
unlike that of the two last-named species, includes a substantial number of insects. 

The parallelism between Hoplotilapia and Macropleurodus extends to the method 
of feeding, and particularly the manner in which the hard-shelled prey is crushed. It 
is this characteristic which most clearly emphasizes the morphological disparity 
between the two genera under discussion on the one hand and the three mollusc- 
eating species on the other ; these latter species, although including a monotypic 
genus, have deviated less markedly from the generalized Haplochromis anatomy. 
Whereas in Astatoreochromis alluaudi, H. ishmaeli and H. pharyngomylus the food 
is crushed entirely by means of the hypertrophied pharyngeal bones and teeth, in 
Hoplotilapia and Macropleurodus the food is broken mainly by the peculiarly 
developed jaws and oral dentition, although the relatively poorly-developed 
pharyngeal mill doubtless continues the process. 

Despite functional similarity in the jaws and dentition of Hoplotilapia and 
Macropleurodus, there is considerable divergence in the detailed morphology of these 
elements. On the one hand, in Macropleurodus the jaws are short and stout, with a 
narrow gape ; associated with the stout supporting skeleton, the teeth are strong. 
On the other hand, the dentary and premaxilla of Hoplotilapia, although broad and 
encompassing a wide gape, appear relatively fragile. The dentition of Hoplotilapia, 
when compared with Macropleurodus, is seen to be composed of small and slender 
teeth which are arranged in bands broader both anteriorly and posteriorly than the 
corresponding teeth of Macropleurodus. Since the shells of Gastropoda in Lake 
Victoria are stouter than those of Lamellibranchiata, it would not be unreasonable 
to suppose that the strong, laterally concentrated and enlarged teeth of Macro- 
pleurodus, and the uniformly multi-seriate, finely-pointed teeth of Hoplotilapia, are 


associated with the predominantly gastropod diet of the former species, and the 
predominantly lamellibranch diet of the latter. 

Study material and distribution records 

Museum and Reg. No. 

S.L. (mm.) 



British Museum (Nat. Hist.) : 

1906. 5. 30. 417-418 

. 90 and 106 

. Buganga (Uganda) . 


1909.5.4. 16 


Sesse Is. (Uganda) 




Jinja, Ripon falls 




Lake Victoria 


Genoa Museum : 

C.E. 12.995 (holotype of P. polyodon) 135 



C.E.12.994 .... 




B.M. (N.H.) : 

1955.2. 10. 141, 147-149. 


. Rusinga Is. (Kenya) . 


1955.2. 10. 145 


. Homa Bay (Kenya) . 


1955.2. 10. 142-144, 146. 


. Kamaringa (Kenya) . 


1955.2. 10. 137-140 


Kisumu (Kenya) 




Jinja (Uganda) 


1955.2. 10. 177-178 


. Beach nr. Nasu Point. 


(Buvuma Channel, Uganda) 

1955.2. 10. 170-171 

. 105 and 107 

Pilkington Bay 



1955.2. 10. 176 


Ramafuta Is. 


1955.2. 10. 168-169 


. Yempita Is. (Rose- . 
bery Channel, Uganda) 

y > 

1955.2. 10. 124-132 


Harbour, Entebbe 


1955.2. 10. 133 


Bugonga beach, 


1955.2. 10. 172-175 


Old Bukakata 



1955.2. 10. 150-163 


. Busongwe (Kagera . 
R. mouth, Uganda) 


1955.2. 10. 179 


Beach south of 
Bukoba (Tanganyika) 


1955.2. 10. 164-167 


Majita beach 


1955.2. 10. 134 


Harbour, Mwanza 


1955.2. 10. 135-136 

. 74 and 139 

. Capri Bay, Mwanza . 


Graham (1929) lists the distribution of H. retrodens as follows : 

Kenya Colony : Mbita passage ; Kavirondo Gulf ; Kadimu Bay. 
Tanganyika Territory : Mussonya Bay (Ukerewe Is.); trawl near Bukoba. 



Cichlid fishes of the Haplochromis group, but differing from that genus in having 
the anterior teeth in both jaws procurrent and disproportionately longer than the 
adjacent lateral teeth. Jaws narrowing at the symphysis ; lips thickened. Known 
only from Lake Victoria. 

Type species : Paralabidochromis victor iae sp. nov. 

The single specimen of Paralabidochromis available provides an interesting taxo- 
nomic and phylogenetic problem. No characters have been found which will dis- 
tinguish this fish generically from specimens of the genus Labidochromis Trewavas ; 
a genus otherwise known only from Lake Nyasa. Unfortunately comparisons must 
be limited to characters apparent in preserved material and then only to the few 
specimens available. Nothing is known of the coloration in life of adult males in 
either genus. This is regrettable since coloration might well provide a reliable 
indication of the affinities of the two genera, both in relation to one another, and to 
the species flocks of Victoria and Nyasa (vide Regan, 1921, 686). The presence of a 
dark sub-marginal band on the dorsal fin of Labidochromis vellicans, in contradistinc- 
tion to its absence in Paralabidochromis is probably of some importance. A sub- 
marginal band is not known in any Lake Victoria Haplochromis species, but is 
present in most species of the group of Nyasa genera to which L. vellicans is 
apparently related (vide Trewavas, 1935, p. 71). 

Although on purely morphological grounds it might seem advisable to include 
the Lake Victoria species within the genus Labidochromis, such a decision would 
imply phyletic relationships between the Victoria and Nyasa species closer than 
those between either species and others of its own lake. To avoid this I have given 
greater weight to the difference in colour-pattern than would perhaps have been 
justified if both inhabited the same lake. 

Apart from the presence of the pan- African genera Tilapia and Haplochromis, 
there is no obvious relationship between the Cichlidae of Lakes Nyasa and Victoria. 
Superficial resemblances between individual species, or genera, in the two lakes have 
been associated with differences which point to their being examples of convergent 
evolution (Regan, 1922, p. 159), although it would perhaps be preferable to consider 
this convergence of morphological characters as parallel evolution since the phenome- 
non occurs between species within a group of related genera. 

It would seem most probable, therefore, that Paralabidochromis represents a 
remarkable example of exact and detailed parallel evolution with Labidochromis. 
Apart from the enlarged anterior teeth, neither Labidochromis nor Paralabidochromis 
departs greatly from the generalized Haplochromis type, as represented in the rivers 
of East and Central Africa. Thus, it is possible that the two genera were independ- 
ently evolved from different parental Haplochromis species, which, however, shared 
the generalized facies of fluviatile species. 


Paralabidochromis victoriae sp. nov. 

Fig. io. 


Depth of body 33-0 ; length of head 31-6 per cent of standard length. Dorsal 
profile of head and snout slightly curved and sloping moderately steeply. Preorbital 
depth 167 per cent of head length ; interorbital width 25-0, snout length 33-4, 
diameter of eye 29-2 and depth of cheek 20-8 per cent of head length. Caudal 
peduncle 1-33 times as long as deep, its length 15-8 per cent of standard length. 

I cm. 
Fig. 10. — Paralabidochromis victoriae, <$, holotype. Drawn by Miss L. Buswell. 

Mouth almost horizontal, lips well developed. Posterior tip of the premaxilla 
extending to a point nearer the nostril than the anterior orbital margin. The angle 
between the rami of each jaw is acute giving a beak-like appearance, which is 
enhanced by the peculiar dentition. 

Dentition. Thirty-eight teeth in the outer series of the premaxilla ; the six anterior 
teeth in both jaws procurrent, greatly elongate, slender, slightly recurved and dagger- 
like ; movable (Fig. 10). Postero-lateral outer teeth small and unicuspid, becoming 
progressively larger and weakly bicuspid laterally. Inner series in both jaws com- 
posed of tricuspid and compressed teeth, of which there are three rows in the upper 
and four in the lower jaw ; outermost row of inner teeth in both jaws, somewhat 
enlarged and standing slightly apart from the remaining rows. 

Lower pharyngeal bone triangular, the teeth small and cuspidate, with only the 
median series slightly enlarged. 


Gill rakers short ; seven on lower limb of first arch. 

Scales ctenoid ; lateral line interrupted, with 32 scales. Cheek with 2 scries of 
imbricating scales ; 6 scales between origin of the dorsal fin and the upper lateral 
line ; 7 between pectoral and pelvic fins. Nuchal scales not exceptionally small. 

Fins. Dorsal, XV 8 ; anal, III 9. Pectoral fin 25-0% of standard length ; pelvics 
with the first soft ray produced and extending posteriorly to the third anal spine. 
Caudal truncate, scaled on proximal half only. 

Coloration. Preserved specimen light brown, with indications of seven faint 
transverse bands on the flanks, and a dark lachrymal stripe. Pelvic fins black, all 
other fins colourless. 

Type locality. Sandy littoral, near Nasu Point, Buvuma Channel. 

Described from a single specimen, a male 76-0 mm. S.L. (91-0 mm. T.L.), collected 
by the author whilst seine-netting at night, 29th May, 1951. 


Paralabidochromis victoriae is closely related morphologically, to Labidochromis 
vellicans of Lake Nyasa, from which it differs principally in possessing large scales 
on the chest, cheek and nape. The dorsal profile of the head and snout in L. vellicans 
apparently differs from that of P. victoriae, in being straighter and more steeply 

Amongst the Haplochromis species of Lake Victoria, P. victoriae, has some structural 
affinity with Haplochromis chilotes. Both species have short and narrow jaws, 
thickened lips and a certain similarity of dentition. For example, the teeth of H. 
chilotes are slender and elongate anteriorly, few in number and arranged in an acute 
dental arch. However, the disproportionately long and procurrent anterior teeth of 
P. victoriae, and the hypertrophied lips of H. chilotes, immediately serve as diagnostic 
characters. The phylogeny of H. chilotes, is at present, obscure, but the species 
would seem more closely related to the H. crassilabris species group in Lake Victoria, 
than to Paralabidochromis. 


It is clear from Regan's analysis of the Lake Victoria Cichlidae, that he did not 
consider the monotypic genera far removed phylogenetically from certain Haplo- 
chromis species, a view which is strengthened by the additional data now available. 
Although the monotypic genera are readily defined by trenchant characters they 
retain fundamental affinities with the Haplochromis species of Lake Victoria. But 
the morphological differentiation which these genera have undergone creates an 
impression of greater divergence than is shown by other adaptive groups within the 
Lake Victoria species flock. Analysis of the diagnostic characters of the genera 
described here, shows that, in any one genus, the anatomical characters of the head 
are functionally related to the dentition. Further, the ontogenetic basis for most of 
these characters is probably attributable to differential growth and not to any 
large-scale qualitative change. Thus the first evolutionary steps involved may well 
have been relatively simple and similar, in the earliest stages, to those which initiated 
the often slight differences characterizing the greater number of Haplochromis species. 


Despite fundamental inter-relationship, three of the four monotypic genera 
exhibit considerable inter-generic divergence in the characters affected by the peculiar 
development of their dentition. The degree of divergence and the means by which 
it has been effected in Macropleurodus, Platytaeniodus and Hoplotilapia are such 
that the genera cannot be related inter se. That is to say, these genera represent 
three independent evolutionary offshoots from the basic Haplochromis stock. The 
fourth genus, Paralabidochromis, shows least departure from the generalized Haplo 
chromis condition, but must also be looked upon as having evolved independently. 

A broad outline of the probable phyletic relationships within the Haplochromis 
species flock is perhaps necessary before attempting further discussion of the 
monotypic genera. 

There are some Lake Victoria Haplochromis species which, when seen in isolation 
or when known only from a few specimens, might seem almost as distinctive as the 
monotypic genera. It is possible, however, to relate these outstanding species to 
others more typical of their particular adaptive group (mollusc eaters, piscivores, 
epiphytic-algal grazers, etc.). Furthermore, when larger series of specimens are 
examined, infra-specific variation is sufficiently high to reduce considerably any 
apparent inter-specific gap. 

With few exceptions, specific differences amongst the Haplochromis are quantitative 
and generally consequent upon the differential growth of various syncranial parts. 
Qualitative differences, on the other hand, are usually those which can be related to, 
and are used in defining, adaptational groups. At both inter- and intra-group levels, 
qualitative and quantitative differences tend to be small and intergrading. 

It is in both qualitative and quantitative characters that the monotypic genera 
depart most markedly from Haplochromis but, unlike the inter-group differences 
within the Haplochromis flock, the morphological gap is clear cut, and remains so 
even when large series of specimens are examined. 

On this interpretation, Hoplotilapia and Platytaeniodus probably examplify the 
phenomenon of " quantum evolution " as described by Simpson (1944 and 1953). It 
is difficult to suggest any ancestry for these genera nearer than a present-day species 
group, viz. the H. crassilabris complex. Even at this level relationship is extremely 
tenuous, and based only on similarity of skull form, in itself probably an adaptational 
character and therefore of doubtful phylogenetic significance. The jaws and dental 
pattern are so dissimilar in Hoplotilapia and Platytaeniodus that one must consider 
the genera separately and not as elements of a single lineage. 

Macropleurodus is less readily regarded as being a product of quantum evolution, 
and will be discussed later. 

Simpson (1944) has noted that quantum evolution is usually associated with a 
shift from one adaptive zone to another, and that the interzonal populations or 
species would be relatively ill-adapted, unstable and short-lived. Thus, morpho- 
logical discontinuity is generally observed between the parental and divergent 

The morphological discontinuity existing between Platytaeniodus and Hoplotilapia, 
and between these two genera and Haplochromis has been demonstrated ; it is the 
more regrettable then that pertinent ecological data for P. degeni are both inadequate 


and confusing, so that it is difficult to equate the apparent morphological specializa- 
tion of this species with any particular ecological niche. The food of P. degeni is 
virtually unknown, although inorganic material and scant but varied organic 
remains suggest bottom-feeding habits. A multi-serial and concentrated dentition, 
such as that of P. degeni may possibly have adaptive value, particularly if food must 
be freed from the substrate or if it requires trituration before digestion can be effected. 
(It should be noted that no Haplochromis has consistently yielded such baffling 
residua in the gut.) 

Hoplotilapia retrodens is somewhat better understood. In this species food is 
predominantly Molluscan. The functional significance of the jaw structure and 
dental pattern in Hoplotilapia, particularly with regard to its observed diet of thin- 
shelled bivalves, has been discussed above (p. 325). From these characters and the 
predominance of Mollusca over the other food organisms it would seem that the 
essential elements of quantum evolution are fairly well defined in H. retrodens. That 
is to say, the species shows both morphological discontinuity and entry into an 
adaptive niche different from that of the presumed parental stock. 

There is, of course, the possibility that Platytaeniodus and Hoplotilapia represent 
an early stage in quantum evolution and that these species may be relatively ill- 
adapted to existing ecological conditions. Their further evolutionary development 
or, alternatively, extinction, will therefore depend upon environment change effecting, 
or failing to effect, the realization of characters which at present could only be 
considered prospective adaptations (sensu Simpson, 1953, p. 188). 

The slight and often indeterminable adaptive differences between Haplochromis 
species occupying the same ecological zone, together with numerous instances of 
inter-specific overlap in feeding habits, would seem to suggest that there is, and has 
been, only slight selection-pressure acting through competition for food. Thus the 
continued existence of Platytaeniodus and Hoplotilapia, like that of many closely 
related Haplochromis species, could be attributed to a period of decreased selection 

Although Macropleurodus, like Platytaeniodus and Hoplotilapia, is separated from 
Haplochromis by a clearly defined morphological gap, the gap is of lesser degree. 
Whereas P. degeni and H. retrodens exhibit unique jaw morphology and dental 
patterns without departure from Haplochromis in tooth form, the basic dental 
pattern of M. bicolor is foreshadowed in two Haplochromis species, as is the form of 
the dentary. Only in tooth form and shape of the premaxilla does M. bicolor show 
great differentiation from H. prodromus. The latter species is morphologically 
closely related to H. sauvagei, a smaller species, which in turn shows departure from 
the basic Haplochromis type towards H. prodromus. In H. sauvagei and H. prodromus 
the outer teeth are relatively stout and, anteriorly, there are several rows of inner 
teeth ; the neurocranium and dentary of these species also approach the M. bicolor 
condition. Both species feed on Mollusca and Insect a, the very slight differences in 
the feeding habits of H. prodromus and H . sauvagei being attributable to the smaller 
size of H. sauvagei. 

It is tempting, therefore, to consider the members of the series H. sauvagei- 
H. prodromus-M. bicolor as representing stages of a lineage, although the possibility 


of independent evolution of the same adaptive characters, cannot be ignored. Until 
more critical evidence is available for the phyletic relationship of these species, they 
may be regarded either as separate end points of different lineages, or as a " stufen- 
reine", or single phyletic line. The three species do illustrate the gross anatomical 
and functional stages through which Macropleurodus could have evolved, although 
the transition in shape of teeth from a typical Haplochromis to Macropleurodus is 
not represented in any living species. 

The isolated position of Paralabidochromis in relation to the Haplochromis species 
of Lake Victoria was commented upon above. 

Little information is available on the evolutionary relationships of Paralabido- 
chromis. Although taxonomically isolated from Haplochromis the genus is nearer 
the Haplochromis stem than either Hoplotilapia, Platytaeniodus or Macropleurodus. 
In Lake Nyasa, Trewavas (1935 and 1949) considers Labidochromis vellicans as 
belonging to a group of nine genera (which group excludes Haplochromis), that, 
although lacking an absolute character to distinguish them, are more closely related 
to each other than to any other genus. Such grouping is impossible for the monotypic 
genera of Lake Victoria. Within this lake the divergent genera of the Haplochromis 
group must be considered as being distinct from one another, as well as from the 
parental stock. 


(1) The genera Macropleurodus Regan, 1922, Platytaeniodus Boulenger, 1906, and 
Hoplotilapia Hilgendorf, 1888, are redefined on the basis of new and fairly extensive 
collections. Similarly, the species M. bicolor (Blgr.) 1906, P. degeni Boulenger, 1906, 
and H. retrodens Hilgendorf, 1888, are redescribed. 

(2) Generic and specific characters are discussed, with particular regard to onto- 
genetic changes. Information gained from small specimens shows that three species 
previously considered as Haplochromis must be added to the synonymies of M. 
bicolor and P. degeni. 

(3) Comparative anatomical and osteological studies of the head indicate that 
Hoplotilapia and Platytaeniodus are not closely related to any extant Haplochromis 
species in Lake Victoria. Morphological stages leading to the syncranial type found 
in Macropleurodus are, however, represented by two endemic Haplochromis species. 
On the basis of syncranial morphology, it is clear that the monotypic genera are not 
closely related inter se. 

(4) Apparently sex-limited polychromatism occurring in Macropleurodus and 
Hoplotilapia is described and discussed. 

(5) A fifth monotypic genus, Paralabidochromis victoriae is described. This genus 
exhibits remarkably close morphological parallelism with Labidochromis vellicans 
from Lake Nyasa. 

(6) Locality lists and notes on the ecology of the genera are given, together with 
observations on the feeding habits of Macropleurodus and Hoplotilapia. 

(7) The evolutionary relationships of the four genera are discussed. It is suggested 
that Hoplotilapia and Platytaeniodus may represent the products of low-level 
quantum evolution. 


A C K N O W L E D G M E N T S 

I wish to acknowledge my gratitude and thanks to the Trustees of the British 
Museum (Natural History) for facilities afforded me during the tenure of a Colonial 
Fisheries Research Studentship ; to Professor L. Bertin of the Museum National 
d'Histoire naturelle, Paris, for allowing me to study type specimens of Lake Victoria 
Cichlidae described by Pellegrin ; to Dr. Delfa Guiglia of the Museo Civico di Storia 
Naturale, Genoa, for the courtesies mentioned in the text ; to Dr. Kurt Deckert, 
Zoologisches Museum, Berlin, for attempting to locate the type specimen of Hoplo- 
tilapia retrodens ; to the officers of the Lake Victoria Fisheries Service who collected 
much valuable material ; to my colleagues Miss R. H. Lowe and Dr. P. S. Corbet 
for many discussions and to the latter for his invaluable identifications of material 
from gut-contents ; and to Mr. Denys W. Tucker, for his very helpful criticism of 
the manuscript. I am especially indebted to Dr. Ethelwynn Trewavas for the 
interest she has shown in my work, and for much helpful information and advice. 


This list includes only those references which occur in the general sections of the text. All 
others appear under the corresponding species headings: 

Cott, H. B. 1952. In East African Fisheries Research Organization Annual Report. East 
Africa High Commission, Nairobi (p. 21). 

Graham, M. 1929. A report on the Fishing Survey of Lake Victoria, 1927-1928, and appendices. 
Crown Agents, London. 

Greenwood, P. H. 1954. On two cichlid fishes from the Malagarazi river (Tanganyika), 
with notes on the pharyngeal apophysis in species of the Haplochrotnis group. Ann. Mag. 
nat. Hist. (12) 7 : 401. 

Regan, C. T. 1920. The classification of the fishes of the family Cichlidae. I. The Tangan- 
yika genera. Ann. Mag. nat. Hist. (9) 5 : 33. 

■ — — 192 1. The cichlid fishes of Lake Nyasa. Proc. zool. Soc. Lond. 675. 

— ■ — 1922a. The cichlid fishes of Lake Victoria. Proc. zool. Soc. Lond. 157. 

1922&. The classification of the fishes of the family Cichlidae. II. On African and 

Syrian genera not restricted to the Great Lakes. Ann. Mag. nat. Hist. (9) 10 : 249. 

Simpson, G. G. 1944. Tempo and Mode in Evolution. New York. 

1953- The Major Features of Evolution. New York. 

Trewavas, E. 1933. Scientific results of the Cambridge expedition to the East African lakes, 

1930-1. II. The cichlid fishes. /. linn. Soc. {Zool.) 38 : 309. 

T 935- A synopsis of the cichlid fishes of Lake Nyasa. Ann. Mag. nat. Hist. (10) 16 : 65. 

— — 1949. The origin and evolution of the cichlid fishes of the great African lakes, with 

special reference to Lake Nyasa. Rapp. 13 e Congres Intern. Zool., Section 5B, 365. Paris. 


- Z MAH mb 



M. SARS, 1846 





ZOOLOGY Vol. 3 No. 8 

LONDON: 1956 





Pp. 335-362 ; Pis. 11-12 ; 2 Text-figures 



ZOOLOGY Vol. 3 No. 8 

LONDON: 1956 

(NATURAL HISTORY), instituted in 1949, is 
issued in five series, corresponding to the Departments 
of the Museum, and an Historical Series. 

Parts will appear at irregular intervals as they 
become ready. Volumes will contain about three or 
four hundred pages, and will not necessarily be com- 
pleted within one calendar year. 

This paper is Vol. 3, No. 8 of the Zoological Series. 


Issued February, 1956 Price Ten Shillings. 



M. SARS, 1846 



During the past century nearly forty hydroids have been referred at one time or another 
to the genus Perigonimus M. Sars. The type species was demonstrated to be a Bougainvillia 
in 1938, and the status of all the other " Perigonimus " spp. is now considered and their position 
determined in accord with modern ideas on classification. They are distributed between the 
Orders Anthomedusae (Athecata), Leptomedusae (Thecata) and Limnomedusae, the majority 
being referred to four families of the first-mentioned Order. 

Some material of Perigonimus vestitus, determined by Allman, has been re-discovered in the 
British Museum and is found to belong to Leuckartiara octona (Fleming) . The original drawings 
of Joshua Alder's Perigonimus linearis, also in the Museum, reveal the presence of oral tentacles 
on the manubrium of the newly-liberated medusa, thus confirming that Bougainvillia flavida 
Hartlaub is identical with it. 


In 1938, I published a partial revision of Perigonimus in so far as it affected the 
groundwork for a monograph of British medusae, later published by F. S. Russell 
( I 953)- The earlier revision facilitated the establishment of common names for 
both the hydroid and the medusa of British species, but did not indicate the position 
of many extra-territorial ones whose status needed to be clarified. In this revision 
the position of these remaining species is considered and the conclusions reached 
earlier for others are re-stated briefly. 

Michael Sars (1846) described a new handsome hydroid from the Norwegian 
coast under the name Perigonimus muscoides and also gave a brief, but inadequate, 
description of the medusa which is liberated with four tentacles. Later Allman 
(1864a), in his revision of Bougainvillid and other hydroids, referred a number of 
them which could not be placed in Bougainvillia (because they did not produce 
Bougainvillid medusae) to this genus of Sars on the assumption that they were 
cogeneric. Thus Perigonimus came to represent a group of Bougainvillia-like 
hydroids, which, either did not give rise to Bougainvillia medusae or had fixed 
gonophores. Gradually the picture emerged that the medusae of most of these 
non-Bougainvillia species belonged to another family, the Pandeidae. Hartlaub 
(1897 and 1914) demonstrated that Perigonimus repens was the hydroid of " Tiara 
pileata", now known as Leuckartiara octona (Fleming), and that another Pandeid, 
Halitholus cirratus Hartlaub, also had a Perigonimus hydroid. More recently Rees 
& Russell (1937) were able to link the Perigonimus serpens kinds of hydroids with 
two species of medusae also of this family. Further revision of Perigonimus was 

zool. 3, 8, 


made possible by the discovery that the type species of Perigonimus gave rise to a 
Bougainvillia medusa (Rees, 1938). Consequently Perigonimus became a synonym 
of Bougainvillia and no longer available as a name for hydroids which could not be 
placed in the same genus or even the same family. 

As Amphinema (i.e., Stomotoca) had already been established by Rees & Russell 
(1937) for the hydroids as well as the medusae of this genus, Leuckartiara (already 
re-established for the medusa phase) was adopted for the appropriate hydroids, and 
Rhizorhagium M. Sars re-established for non-medusa bearing species. 

As already mentioned this left a number of " Perigonimus " species either not 
satisfactorily classified or left out of consideration, and, in the brief notes below, 
it has been thought desirable to indicate their present status and where possible to 
add new information. 


Perigonimus abyssi G. O. Sars 
Perigonimus abyssi G.O. Sars, 1873, Forh.Vidensk. Selsk. Krist., 1873 : 126, Tab. v, figs. 27-30. 
The newly liberated medusa of this minute species becomes free with two opposite 
tentacles and two partly developed tentacles on the other perradial bulbs (Rees, 
1938). It is a Pandeid and has been referred provisionally to Leuckartiara Hartlaub 
pending further knowledge of its possible relationship with medusae of the genera 
Leuckartiara, Neoturris or Catablema. 

Perigonimus antarcticus Hickson & Gravely 

Perigonimus antarcticus Hickson & Gravely, 1907, Nat. Antarct. Exped., 1901-1904, Nat. Hist., 
3 : 1, pi. i, figs. 1-3 ; pi. iv, fig. 32. 

This species was placed in Atractylis (synonymous with Bougainvillia) by Van- 
hoffen (1910) and in a new genus Gravely a by Totton (1930), but its affinities lie 
with Rhizorhagium M. Sars, to which genus it was referred by Stechow (1919) and 
by Rees (1938). This last mentioned genus seems to be well constituted for simple 
Bougainvillid hydroids with fixed gonophores, but its relationship to Aselomaris 
Berrill and Garveia Wright will have to be considered in a wider revision of the 

Perigonimus apicatus Brooks 
Perigonimus (Stomotoca) apicata Brooks, 1884, Zool. Anz., 7 : 711. 

This is the name given by Brooks to the hydroid of Stomotoca apicata, now known 
as Stomotoca dinema (Peron & Lesueur). Mayer (1910, p. 113) referred this hydroid 
to 5. rugosa Mayer, but the absence of an apical projection in the newly liberated 
medusa points to 5. dinema, as does the name used by Brooks for his hydroid. 

Perigonimus bitentaculatus (Wright) 

Atractylis bitentaculata Wright, 1867, Proc. R. Phys. Soc. Edinb., 3 : 45, pi. i, fig. 5. 

This species was referred provisionally to Perigonimus by Hincks (1868), but 
Wright himself gives the best clue as to the relationships of his species in the following 


sentence : " They have the habit, like that of Lar (Gosse), of quickly and repeatedly 
bending down the body until the mouth is brought close to the surface on which 
the zoophyte grows." 

This species has not been reported since it was found by Wright in a Pecten shell 
dredged from the Firth of Forth near Inch Keith. The characters of the hydranth 
with its two opposite tentacles strongly suggest that the hydroid is related to Pro- 
boscidactyla (Lar of Gosse) and should be placed in the same family. The possibility 
should not be overlooked that this species may prove to belong to Pochella polynema 
Hartlaub, the only other medusa of this family known to occur in British waters. 
I do not propose to create a new genus for this species, but merely note that its 
position appears to be in the family Proboscidactylidae of the Order Limnomedusae. 

Perigonimus cidaritis Weismann 

Perigonimus cidaritis Weismann, 1883, Entstehung der Sexualzellen bei den Hydromedusen, Jena, 
p. 117, pi. xii, fig. 10— 11. 

This hydroid has a well-marked pseudohydrotheca in which the bases of the 
tentacles are protected in tube-like extensions of the perisarc as in Bimeria and 
Thamnostoma. As the medusa is a Bougainvillid closely related to Thamnostoma 
russelli Rees and there is very little difference in the hydroids, P. cidaritis has been 
referred to Thamnostoma by Rees (1938). 

Perigonimus cirratus Hartlaub 
Perigonimus cirratus Hartlaub, 1914, Nord. Plankt., Lief. 17, XII : 274, fig. 234. 

Hartlaub (1914) gave the name P. cirratus to the hydroid of the medusa Halitholits 
cirratus which he described in the same memoir, presumably for the reason that it 
was then customary to adhere to a dual system of classification. There are now no 
legitimate reasons why the hydroid should not also take the name Halitholus. 

Perigonimus coccineus (Wright) 
Atractylis coccinea Wright, 1861, Ann. Mag. Nat. Hist., (3) 8 : 130. 

E. S. Russell (1907) is the only author who has reported this species since it was 
originally described by Wright. In 1938 I suggested that the species might prove to 
be identical with Rhizorhagium roseum. 

Perigonimus confertus (Alder) 
Eudendrium ? confertum Alder, 1856, Trans. Tyneside Nat. Fid. CI., 3 : 103, pi. iii, figs. 5-S. 
Allman (1859) placed this species in Dicoryne because of the peculiarities of the 
gonophores, an assessment which has been generally approved. Recently Vervoort 
(1946) used the name Perigonimus confertus for the first time for this species, although 
presumably aware that this name is synonymous with Bougainvillia and therefore 
no longer available. The name Dicoryne conferta has been the established name of this 
species since 1859 and I see no reason to change it. 


Perigonimus decorans Schneider 
Perigonimus decorans Schneider, 1897, Zool. Jb. Syst., 10 : 479. 

Schneider's description is too inadequate even to indicate in what family this 
hydroid could be placed. 

Perigonimus formosus Fewkes 
Perigonimus formosus Fewkes, 1889, Bull. Essex Inst., 21 : 102, no figure. 

Some small thumb-nail sketches of this species have been given by Fraser (1937, 
p. 35, pi. v, fig. 20) ; they indicate that P. formosus should be placed in Rhizorhagium 
M. Sars. A new description, especially of the gonophores, is much needed to determine 
the relationship of the species to R. roseum. 

Perigonimus gelatinosus Duerden 

Perigonimus gelatinosus Duerden, 1895, Sci. Proc. R. Dublin Soc, n.s., 8 : 325, 327, pi. xiv, 
figs. 2-3. 

Duerden's species is not identifiable with certainty with any known British species, 
although there is a possibility that it is identical with Leuckartiara octona as supposed 
by Hartlaub (1914). The complete retraction of the hydranth within the pseudo- 
hydrotheca could be due to poor preservation and by itself is no indication of specific 

Perigonimus georginae Hadzi 

Perigonimus georginae Hadzi, 1913, Rad. Jug. Akad. Znan. Umj., 200 : 98, figs. 7, 10 and 11. 

The original description by Hadzi is not very satisfactory. He appears to have 
confused at least two species. His figure 7 is a Stomatoca hydroid, his figure 8 repre- 
sents a Bougainvillia, and the newly liberated medusa, although combining some of 
the features of two species of Stomotoca medusae, cannot be referred with certainty 
to either 5. rugosa Mayer or to 5. dinema (Peron & Lesueur). Until the Adriatic 
medusae have been re-investigated it seems preferable to let this species retain its 
identity as Stomotoca georginae. The name Dinema applied by Hadzi to the medusa 
is a synonym of Leuckartiara, a member of a different sub-family. 

Perigonimus (?) inflatus Duerden 
Perigonimus (?) inflatus Duerden, 1895, 5a. Proc. R. Dublin Soc, n.s., 8 : 329, pi. xiv, fig. 4. 
This little known species does not seem to have been reported since Duerden's 
original description. It is here referred provisionally to Rhizorhagium M. Sars. 

Perigonimus jonesii Osborn & Hargitt 
Perigonimus jonesii Osborn & Hargitt, 1894, Amer. Nat., 28 : 27, figs. 1-12. 

I have followed Hartlaub (1914) in placing this species in the synonymy of 
Leuckartiara octona (Fleming). The colony described by Osborn & Hargitt compares 
favourably with those found on Scaphander lignarius in British waters. 


Perigonimus linearis (Alder) 

Atractylis linearis Alder, 1862, Trans. Tyneside Nat. Fid. CI., 5 : 230, pi. x, figs. 1-3. 

Alder's Atractylis linearis was tentatively referred to Perigonimus byAllman (1864a). 
The newly liberated medusa has a very thick jelly and a small subumbrellar cavity, 
and in this it resembles Bougainvillia flavicla Hartlaub (1897), an opinion later 
expressed by Hartlaub himself (1911). The published drawings of Alder do not 
suggest any oral tentacles on the manubrium of the medusa, but fortunately his 
original drawings (from which the engraving for his paper (1862, pi. x, figs. 2 and 3) 
was prepared) are preserved in the British Museum (Nat. Hist.), having been acquired 
by Canon A. M. Norman and later bequeathed by him. These drawings do show the 
oral tentacles faintly and so the two species can be regarded as identical. 

Russell (1953) does not venture an opinion on the status of P. linearis or its medusa, 
but suggests that B.flavida may prove to be the hydroid of Bougainvillia britannica. 
Until this relationship can be demonstrated I propose to retain Alder's species under 
the name Bougainvillia linearis (Alder, 1862). 

Perigonimus maclovianus (Lesson) 

Perigonimus maclovianus Vanhoffen, 1910, Dtsch. Sudpolar-Exped., 11 : 272, fig. loa-d (hydroid). 
The medusa of this species was described under the name Cyanaea bougainvillii 
by Lesson (1830), who later made it the type species of Bougainvillia under the name 
B. macloviana (1836). This is the name in common use to-day for both hydroid and 
medusa, although Vanhoffen (1910), who first described the hydroid, referred it to 
Perigonimus. Both Stechow (1919, p. 21) and Kramp (1928, p. 50) pointed out that 
Bougainvillia was the correct generic name for both hydroid and medusa. 

Perigonimus minutus Allman 
Perigonimus minutus Allman, 1863, Ann. Mag. Nat. Hist., (3), 11 : 11. 

Allman's Perigonimus minutus has been referred to Leuckartiara oclona both by 
Hartlaub (1914) and Rees (1938). Allman's description and figures (1871-2, pi. xi, 
figs. 4-6) correspond closely with the slender polyps such as are commonly found on 
Scaphander lignarius (Rees, 1938, p. 14 fig. 3d-f). 

Perigonimus miniatus (Wright) 

Atractylis miniata Wright, 1863, Proc. R. Phys. Soc. Edinb., 2 : 351. 

Wright's Atractylis miniata was referred provisionally to Perigonimus by Hincks 
(1868). The reproduction of the species is not known, but the description suggests that 
Wright was describing a colony of Rhizorhagium and R. roseum in particular. 

Perigonimus multicornis Allman 
Perigonimus multicornis Allman, 1874, Nature, Lond., 11 : 179. 

A re-examination of Allman's type material in the Zoological Museum, Copenhagen 
by Kramp (1926) revealed that the species was Eudendrium ramosum. 



Perigonimus muscoides M. Sars 
Perigonimus muscoides M. Sars, 1846, Fauna Littoralis Norvegiae, Heft. 1 : 8, Tab. 1, figs. 19-21. 
This is the type species of Perigonimus and the elucidation of its life history by Rees 
(1938, p. 2) revealed that its medusa was a Bougainvillia, B. nordgaardi (Browne, 
1903). Perigonimus M. Sars, 1846, thus falls into the synonymy of Bougainvillia 
Lesson 1836 and the name Bougainvillia muscoides becomes applicable to both 
hydroid and medusa as demonstrated in the above paper. 

Perigonimus muscus Allman 
Perigonimus muscus Allman, 1863 ; Ann. Mag. Nat. Hist. (3), 11 : 12. 

This species was referred to Bougainvillia by Allman (18640). It was considered to 
be synonymous with Bougainvillia ramosa by Hartlaub (1911) and this view was 
confirmed by the studies of Brink (1925) who demonstrated that colonies passed 
through a muscus stage when young before reaching the stage typified by the usual 
ramosa kind of colony. 

Perigonimus (?) nanellus Stechow 
Perigonimus (?) nanellus Stechow, 1919, Zool. Jb. Syst., 42 : 14, fig. C. 

I consider this species to be a Bougainvillia, possibly representing a well formed 
colony of one of the common species, and pending further knowledge of its life 
history it is now referred to this genus as B. nanellus. 

Perigonimus napolitanus Hargitt 
Perigonimus napolitanus Hargitt, 1904, Mitt. zool. Sta. Neapel, 16 : 571, taf. 22, fig. 25. 
This species is not recognizable. 

Perigonimus nudus Stechow 
Perigonimus nudus Stechow, 1919, Zool. Jb. Syst., 42 : 16, fig. D. 

The form of the hydranth as noted by Stechow resembles those of the medusa- 
bearing Campanopsinae of the family Haleciidae and the relationships of the species 
undoubtedly lie with this group rather than with any Athecate (Anthomedusan) 

Perigonimus ? nutans Hincks 
Perigonimus (?) nutans Hincks, 1877, Ann. Mag. Nat. Hist., (4), 19 : 149, pi. xii, fig. 1. 

Rees & Russell (1937, p. 71) regard this species as possibly a young polyp of the 
Perigonimus serpens type ; this kind of hydroid belongs to the genus Amphinema. 
There are two species in British waters, A. dinema and A. rugosum, and these are so 
alike that it is not possible to assign this species to one or the other. 

Perigonimus palliatus (Wright) 
Atractylis palliata, Wright, 1861, Ann. Mag. Nat. Hist. (3), 8 : 129, pi. iv, figs. 6 and 7. 

In this species the pseudohydrotheca is large and dilated, but I do not think it has 
any special significance as a diagnostic character. The medusa, according to Wright 


(1861) " resemble exactly those of Atractylis (Perigonimus) repens ", that is, Leuckarti- 
ara octona (Fleming). Wright's colony was on a shell inhabited by a hermit crab 
Eupagurus bernhardus and the size of the pseudohydrotheca can be attributed to 
habitat and feeding conditions as in the other colonies of Leuckartiara octona living 
under similar conditions (see Rees, 1938, p. 16, fig. 5). I regard the two species as 

Perigonimus pugetensis Heath 
Perigonimus pugetensis Heath, 19 10, Biol. Bull. Woods Hole, 19 : 74, figs. 1 and 2. 

This small hydroid was found growing on a fish, Hypsagonus quadricornis (C. & V.) ; 
it was found that its newly liberated medusa was a Pandeid like Leuckartiara octona. 

Hartlaub (1914) doubted whether the species was distinct from L. octona, and 
the latter has since proved to be a cosmopolitan species. Rees (1938) found a colony 
of L. octona growing on a fish, Agonus cataphr actus, and this colony compared very 
favourably with that of Heath. It is now proposed to regard P. pugetensis as a syno- 
nym of L. octona. 

Perigonimus pusillus (Wright) 
Eudendrium pusillum Wright, 1857, Edinb. New Phil. J., N.s., 6 : 84, pi. ii, fig. 8, 9. 

Stechow (1919) revived the name pusillus for the better known name Perigonimus 
repens (Wright, 1858). It appears that Wright changed the name of his species to 
repens on the receipt of a paper by M. Sars (1857) in which he described a species of 
Halecium under the name Eudendrium pusillum. 

Wright's Species falls into the synonymy of Leuckartiara octona (Fleming). 

Perigonimus quadritentaculatus (Wright) 
Atractylis quadritentaculata Wright, 1867, Proc. R. Phys. Soc. Edinb. 3 : 45, pi. i, fig. 6. 

This creeping form with long stolons and sessile, four tentacled hydranth is possibly 
a juvenile Bougainvillid, which cannot be identified with certainty with any particular 

f Perigonimus robustus Fraser 
? Perigonimus robustus Fraser, 1938, Rep. Hancock Pacific Exped., 4, No. 1 ; 17, pi. iii, fig. 12. 
In the absence of information on the gonophore this species cannot be satisfactorily 
classified, and is therefore provisionally placed in Bougainvillia. 

Perigonimus roseus (M. Sars) 
Rhizorhagium roseum M. Sars, 1877, Fauna Littoralis Norvegiae, Heft 3 : 28, Tab. ii, figs. 37-43. 
The genus Rhizorhagium was created for this species by Sars, and although the 
type species was referred for a time to Perigonimus by Bonne vie (1899) and others, 
the genus was re-instated by Rees (1938). 

Perigonimus sarsii Bonnevie 

Perigonimus sarsii Bonnevie, 1901, Bergens Mus. Meeresfauna von Bergen, Heft I, p. 6. 

This species has been referred to Rhizorhagium by Rees (1938) and there is a 
distinct possibility that the species is founded on a colony of R. roseum, growing over 
the stems of another hydroid, 

ZOOL. 3, 8, 


Perigonimus schneideri Motz-Kossowska 

Perigonimus schneideri Motz-Kossowska, 1905, Arch, Zool. exp. gen., (4), 3 : 72, fig. VI. 

Stechow (1922) created a new genus Perarella for this species and indicated that 
its position was not in the Hydractinidae but rather in the Bougainvillinae near 

Since then Komai (1931) has described the hydroid of the medusa Cytaeis japonica, 
to which the present species is clearly related. Both species have the following 
features in common : a reticulate anastomozing network of stolons, a short collar 
round the base of the hydranth, the hydranth itself long and tubular, of the Podo- 
coryne type, and the gonophores borne singly on the hydrorhiza. 

I propose to retain Perarella for Cytaeid hydroids in which there are fixed gonophores 
and to place the genus in the family Cytaeidae, L. Agassiz, 1862. Agassiz's definition 
excludes unrelated forms such as Podocoryne (including Lymnorea), Turritopsis and 
Oceania, which Uchida (1927) and others have placed in the family. 

While possessing distinct oral tentacles on the manubrium of the medusa as in the 
Bougainvillidae, the presence of simple perradial tentacles together with the characters 
of the hydroid justify the existence of a separate family for Cytaeis and Perarella — a 
family in which the combined characters of hydroid and medusa present some 
characters in common with both the Hydractinidae and the Bougainvillidae, but 
none of their specialized features. 

Perigonimus serpens Allman 

Perigonimus serpens Allman, 1863, Ann. Mag. Nat. Hist. (3), 11 : 10. 

Rees and Russell (1937) have referred this species to Amphinema dinema following 
the elucidation of the life history of this medusaand its hydroid. Allman (1871-2) clear- 
ly indicated in his illustrations of the medusa that he was dealing with this species 
and not with the closely related A . rugosum. 

Perigonimus sessilis (Wright) 
Eudendrium sessile Wright, 1857, Edinb. New Phil. J, n.s., 6 : 90, pi. iii, figs. 16-17. 

This is the sessile form of Leuckartiara octona (Fleming). When the colony is 
growing on an uneven, constantly abraded surface the hydranths remain sessile in 
protected hollows or grooves and do not progress beyond the sessilis stage (Rees, 

Perigonimus steinachi Jickeli 
Perigonimus steinachi Jickeli, 1883, Gegenbaurs Jb., 8 : 617, pi 27, figs. 1-9. 

Jickeli gives a totally inadequate description of the morphology of his hydroid 
(although he discusses its histology at length) and it is not possible to indicate what 
it might be. 

Perigonimus sulfureus Chun 
Perigonimus sulfureus Chun, 1889, 5. B. Akad. Wiss. Berlin, 1889 (2) : 524. 

Stechow (1921 and 1923) created a new genus Perigonella for this species and placed 
it with the Hydractinians near Stylactella. Steche (1906) figures a well-developed 


medusa bud with conical bulbs with tentacles and manubrium devoid of oral tentacles, 
but we have insufficient information to assign this species to a family. 

Perigonimus vagans Thornely 

Perigonimus vagans Thornley, 1908, /. Linn. Soc. Zool., 31 : 81, pi. ix, fig. I. 

This species is now referred provisionally to Bougainvillia pending the re-discovery 
of the hydroid. Thornely's description is insufficient to permit any other suggestions. 

Perigonimus vestitus Allman 

Perigonimus vestitus Allman, 18646, Ann. Mag. Nat. Hist., Ser. (3), 14 : 57. 

In 1846 Allman described a new Perigonimus, P. vestitus from the Firth of Forth, 
in which, to quote his 1872 diagnosis the " posterior part of the body [of the hydranth], 
invested by the rough perisarc, which is thence continued as a delicate, smooth 
membrane over the remainder of the body nearly as far as the mouth ". His figures 
(1871, pi. xi, figs. 1-3) show the pseudohydrotheca (that is the perisarcal investment 
of the hydranth) to extend anteriorly beyond the tentacles close to the mouth. There 
is no suggestion that this perisarc is perforated to allow the tentacles to project (as 
they do) and no suggestion of any tubular perisarcal sheathing encircling the base of 
any tentacle. This seems to be one of the few occasions on which this far-sighted 
pioneer of hydroid systematics failed to make a satisfactory description. 

Stechow (1919) referred P. vestitus to the medusa genus Cytaeis presumably on an 
assumed relationship between the hydroid and medusa of vestitus with Perigonimus 
cidaritis. There were no real grounds for this, as the figures demonstrate that P. 
vestitus gives rise to a typical Pandeid medusa, so that it must be stated that the 
species is related to genera like Leuckartiara, Amphinema and Halitholus. 

In Perigonimus vestitus too, the bases of the tentacles are not enclosed in perisarcal 
tubes extending from the pseudohydrotheca itself. Such a feature is characteristic 
of Bimeria, Thamnitis and Thamnostoma, and the medusa of the latter clearly has 
affinities with the Bougainvillidae rather than the Pandeidae. Cytaeis, which Stechow 
erroneously associated with these Bimerid hydroids, belongs to none of these families. 

Hartlaub (1914) in his revision of the Tiaridae had listed Perigonimus vestitus 
under the synonymy of Leuckartiara octona, but I was reluctant to follow him in 
my study of variation in the hydroid of this species (1938) in view of the diagnosis 
given by Allman. Fortunately, material of Perigonimus vestitus determined by Allman, 
growing on Lepidochiton cinereus, has recently been found in the collections of the 
British Museum (B.M. No. 1877.4. 12.31). 

A re-examination reveals that although the soft parts have disappeared the 
chitonous perisarc of the stems and pseudohydrothecae are still present. The latter 
are very similar in appearance to those that I described for a colony of Leuckartiara 
octona (Fleming) on Corystes cassivelaunus (Rees, 1938, p. 16, fig. 5). There is no 
suggestion of any perisarcal sheaths such as are found in Bimeria. The colonies 
exhibit no features that could not be found in a well-developed colony of Leuckartiara 
octona, and the characters of the medusa, as described by Allman, support this. For 


these reasons the species vestitus is placed in the synonymy of Leuckartiara octona 

Perigonimus yoldiae-arcticae Birula 

Perigonimus yoldiae-arcticae Birula, 1897, Annu. Mus. Zool. Acad. St. Petersb. 2 : 86, pi. x, fig. 3. 
Jaderholm (1909) gives an excellent figure of this hydroid. I have examined 
material from the following localities : 

Zoologiske Museum, Copenhagen. 

Kandalakscha, on Yoldia arctica, 24 fm. (from St. Petersbourg Museum). 

Riksmuseum, Stockholm. 

Kandalakscha, 21-27 fm. Coll. N. Knipowitsch (from St. Petersbourg Museum). 
Kandalakscha, 20-25 fm. Coll. A. Birula (from St. Petersbourg Museum). 
Nordenskiold's Sea, 77 i 1 N., 114 35 1 E., 3.ix.i90i (Russian Polar Expedition, 

1900-1903, St. 46). 
Nordenskiold's Sea, 75 42 1 N., 124 41 1 E., 6.ix.i90i (Russian Polar Expedition, 

East Greenland, Franz- Josef Fjord (Muskoxfjord), 21.viii.1900, 220 m. (Swedish 

zoological Polar Expedition, 1900, St. 27). 

All the material from the above localities is sterile. The pseudohydrotheca of the 
hydranth reaches almost to the tentacles, and in one hydranth it is a little loose- 
fitting and wrinkled which permits its limits to be seen. There is no indication of a 
web between the tentacles. 

The most interesting (and also the characteristic) feature of this hydroid is the 
formation of ' nodes ' on the stems, and if we accept the belief that the species is a 
gymnoblastic hydroid (as seems most likely), these "nodes" indicate points of 
regeneration after periodic dying down of the hydranths. The species is referred 
provisionally to Bougainvillia. 


Family Cytaeidae 

Genus Perarella Stechow (Original name) 

Perarella schneideri Perigonimus schneideri Motz-Kossowska 

Family Bougainvillidae 

Sub-family Thamnostominae 
Genus Thamnostoma Haeckel 
Thamnostoma cidaritis 
Sub-family Bougainvillinae 
Genus Bougainvillia Lesson 
Bougainvillia macloviana 
Bougainvillia muscoides 
Bougainvillia ramosa 
Bougainvillia linearis 

Perigonimus cidaritis Weismann 

Perigonimus macloviana Vanhoffen 
Perigonimus muscoides M. Sars 
Perigonimus muscus Allman 
Atractylis linearis Alder 


Provisionally also: 

Bougainvillia nanellus 

Bougainvillia robusta 

Bougainvillia vagans 

Bougainvillia yoldiae-arcticae 

Bougainvillia sp. 
Genus Dicoryne Allman 

Dicoryne conferta 
Genus Rhizorhagium M. Sars 

Rhizorhagium roseum 

Rhizorhagium antarcticum 
Rhizorhagium formosum 

Perigonimus (?) nanellus Stechow 
Perigonimus robustus Fraser 
Perigonimus vagans Thornely 
Perigonimus yoldiae-arcticae Birula 
Atractylis quadritentaculata Wright 

Eudendrium confertum Alder 

Rhizorhagium roseum M. Sars 
Perigonimus sarsii Bonnevie 
? A tractylis coccinea Wright 
? A tractylis miniata Wright 
? Perigonimus inflatus Duerden 
Perigonimus antarcticus Hickson & Gravely 
Perigonimus formosus Fewkes 

Family Pandeidae 

Sub-family Stomotocinae 
Genus Stomotoca L. Agassiz 
Stomotoca dinema 

Stomotoca georginae 
Sub-family Pandeinae 

Genus Leuckartiara Hartlaub 
Leuckartiara octona 

Provisionally also : 

Leuckartiara abyssi 
Genus Halitholus Hartlaub 

Halitholus cirratus 

Family Eudendriidae 

Genus Eudendrium Ehrenberg 
Eudendrium ramosum 

Perigonimus serpens Allman 
Perigonimus (?) nutans Hincks 
Perigonimus apicatus Brooks 
Perigonimus georginae Hadzi 

Eudendrium pusillum Wright 

Eudendrium sessile Wright 

Eudendrium repens 

A tractylis palliata Wright 

Perigonimus minutus Allman 

Perigonimus vestitus Allman 

Perigonimus jonesii Osborn & Hargitt 

Perigonimus pugetensis Heath 

? Perigonimus gelatinosus Duerden 

Perigonimus abyssi G. O. Sars 

Perigonimus cirratus Hartlaub 

Perigonimus multicornis Allman 

Family Haleciidae 

Sub-family Campanopsinae 

Perigonimus nudus Stechow 

Family Proboscidactylidae 
Genus Pochella Hartlaub 
? Pochella polynema 


A tractylis bitentaculata Wright 


Incertae Sedis 

Perigonimus decorans Schneider 

Perigonimus napolitanus Hargitt 

Perigonimus steinachi Jickeli 

Perigonella sulphureus [Perigonimus sulphureus Chun) 

These few species which I have been unable to classify are insufficiently described. 

The above classification is in accord with recent progress, but some of it may not 
be familiar to specialists who work only on hydroids and leave the medusa phase out 
of consideration. I believe with Allman (1864a) that "An adequate conception of the 
hydroid can thus only be obtained by regarding it as the product of two factors, one 
of them finding its expression in the trophosome, and the other in the gonosome ". 
The result is frequently a mosaic, a blending of characteristics into a pattern which 
gives a much better picture of the position of the living species than does consideration 
of only a part of the life history. 1 

Much has been written by Kramp (1941 and 1949) about the dangers of attempting 
a single classification for hydroids and medusae, but its outlines are already apparent 
for the Anthomedusae, and this revision of Perigonimus is intended as a contribution 
towards it. 


Agassiz, L. 1862. Contributions to the Natural History of the United States of America, 4 

1-380, pi. xx-xxxiv. 
Alder, J. 1856. A Catalogue of the Zoophytes of Northumberland. Trans. Tyneside Nat. 

Fid. CI., 3 : 93-162 ; pis. iii-x. 

1862. Supplement to a Catalogue of the Zoophytes of Northumberland and Durham: 

Trans. Tyneside Nat. Fid. CI., 5 : 225-247. 

Allman, G. J. 1859. Notes on the British Zoophytes. Ann. Mag. Nat. Hist. (3), 4 : 367- 


1863. Notes on the Hydroida. Ann. Mag. Nat. Hist. (3), 11 : 1-12. 

1864a. On the construction and limitation of genera among the Hydroida. Ann. Mag. 

Nat. Hist. (3), 13 : 345-380. 

18646. Notes on the Hydroida. Ann. Mag. Nat. Hist. (3), 14 : 57-64. 

1871-72. A monograph of the Gymnoblastic or Tubularian Hydroids. xxiv -f- 450 pp. 


1874. On the diagnosis of new genera and species of Hydroids. Nature, Lond., 11 : 179. 

Birula, A. 1897. Recherches sur la biologie et la zoogeographie principalement des mers 

russes. 2. Hydrozoaires, Polychetes et Crustaces recueillis par le Dr. A. Botkine en 1895 
dans les Golfes du Jenisei et de l'Obi. Annu. Mus. zool. Acad. Sci. St.-Petersb., 2 : 78-116. 
Bonnevie, K. 1899. Hydroida. Den Norske N or dhavs -Exp edition 1876-1878, 26 : Zoology, 
104 pp. 

1901. Hydroiden. Bergens Museum Meeresfauna von Bergen, 1 : 1-15. 

Brink, R. 1925. Notes concerning the variability and the action of environmental influences 
on the structure and growth of the hydroid-colony Bougainvillia ramosa (v. Ben.) Lesson 
and its bearings on systematics. Proc. Akad. Sci. Amst., 27 : 726-733. 

Brooks, W. K. 1884. On the life history of Eutima, and on radial and bilateral symmetry 
in Hydroids. Zool. Anz., 7 : 709-711. 

'Basic principles in the classification of hydroids and medusae will be considered in another paper. 


Browne, E. T. 1903. Report on some Medusae from Norway and Spitzbergen. Bergens 

Mus.Avb. No. 4 : 1-36. 
Chun, C. 1889. Beriche uber eine nach den Canarischen Inseln im Winter 1887-88 ausge- 

fuhrte Reise. II Beobachtungen uber die pelagische Tiefsen- und Oberflachenfauna des 

ostlichen Atlantischen Oceans. S-B. preuss. Akad. Wiss., 1889 : 519-553. 
Duerden, J. E. 1895. Notes on the Hydroida and Polyzoa. Ex. Survey of fishing grounds, 

west coast of Ireland 1890-91. Sci. Proc. R. Dublin Soc. (n.s.), 8 : 325-336. 
Fewkes, J. W. 1889. New Invertebrata from the coast of California. Bull. Essex Inst. 21: 

Fraser, C. M. 1937. Hydroids of the Pacific Coast of Canada and the United States. Toronto. 

Pp. 1-207, pl- I-XLIV. 

1938. Hydroids of the 1934 Allen Hancock Pacific Exp. Rep. Hancock Pacific Exped. 

4 : 1-105. 

Hadzi, J. 1913. Poredbena hidroidska istrazivanja II. Perigonimus corii sp. n. i Perigonimus 
georginae sp. n. Rad. Jug. Znan. Umj. Akad. 200 : 89-108, 11 text-figs. 

Hargitt, C. W. 1904. Notes on some Hydromedusae from the Bay of Naples. Mitt. zool. 
Sta. Neapel, 16 : 553-585. 

Hartlaub, C. 1897. Die Hydromedusen Helgolands. Wiss Meeresuntersuch., N.F. Bd. II, 
Abt. Helgoland, Heft. I, X : 449-512. 

191 1. Nord. Plankt. Lief. 15, XII. Craspedote Medusen, Tiel 1, Lief. 2, Familie III, 

Margelidae : 137-236. 

1914. Nord. Plankt. Lief. 17, XII. Craspedote Medusen, Tiel 1, Lief. 3, Familie IV, 

Tiaridae : 237-363. 

Heath, H. 1910. The association of a Fish with a Hydroid. Biol. Bull. Woods Hole, 19 : 

Hickson, S. J. and Gravely, F. H. 1907. Hydroid Zoophytes. Nat. Antarct. Exped. 

1901-1904, Nat. Hist. 3 : 1-34. 
Hincks, T. 1868. A History of the British Hydroid Zoophytes. London : pp. lxviii + 338, 

1 pl. ; Atlas, 67 pis. 

1877. Contributions to the history of the Hydroida. Ann. Mag. Nat. Hist. (4), 19 : 


Jaderholm, E. 1909. Northern and Arctic Invertebrates in the collection of the Swedish 
State Museum (Riksmuseum). IV. Hydroiden. K. Svenska Vetensk. Akad. Handl. 
45 No. 1, 124 pp 

Jickeli, C. F. 1883. Der Bau der Hydroidpolypen. Gegenbaurs Jb. 8 : 580-680. 

Komai, T. 1 931. On the hydroid stage of Cytaeis japonica Uchida. Annot. zool. jap. 13 : 

Kramp, P. L. 1926. Occasional notes on Coelenterata. 1 : a. The type specimens of 
" Perigonimus multicornis " and " Eudendrium rigidum " Allman. b. Eucheilota maculata 
and Campanulina hincksii Hartlaub. c. Cylocanna welshi Bigelow. Vidensk. Medd. 
naturh. Foren. Kbh. 82 : 241-247. 

— ■ — 1928. Papers from Dr. Th. Mortensen's Pacific Expedition 1914-16. XLIII. Hydro- 
medusae, I. Anthomedusae. Vidensk. Medd. naturh. Foren. Kbh., 85 : 27-64, figs. 1-30. 

1941. Notes on the hydroid Campanulina panicula G. O. Sars. Goteborgs Vetensk. 

Samh. Handl. (6) ser. B 1, No. 2 ; 1-11. 

1949- Origin of the hydroid family Corymorphidae. Vidensk. Medd. Dansk. naturh. 

Foren. Kbh. Ill : 183-215. 

Lesson, R. P. 1830. Zoophytes. Voyage autour du monde execute . . . sur La Coquille 
par M. L.-J. Duperry. Zoologie par M. Lesson. 2 vol. Paris 1826-30 (Vol. 2, P. 2, 2e 
division, 1830). 

1836. Memoire sur la familie des Beroides (Beroidae Less.). Ann. sci. nat. {Zool.) (2) 

Vol. 5, pp. 235-266. 

Mayer, A. G. 1910. Medusae of the World. Vol. t. Publ. Carneg. Instn. 109 : 1-230. 


Motz-Kossowska, S. 1905. Contribution a la connaissance des Hydraires de la M6diter- 

ranee occidentale. 1. Hydraires gymnoblastiques. Arch. Zool. exp. gen. (4) 3 : 33-98. 
Osborn, H. L. & Hargitt, C. W. 1894. Perigonimus Jonesii ; a Hydroid supposed to be 

new, from Cold Spring Harbor, Long Island. Amer. Nat. 28 : 27-34. 
Rees, W. J. 1938. Observations on British and Norwegian hydroids and their medusae. 

/. Mar. biol. Ass. U.K., 23 : 1-42. 
Rees, W. J. & Russell, F. S. 1937. On rearing the hydroids of certain medusae, with an 

account of the methods used. /. Mar. biol. Ass. U.K. 22 : 61-82. 
Russell, E. S. 1907. The Atractylis coccinea of T. S. Wright. Ann. Mag. Nat. Hist. (7), 

20 : 52-55- 
Russell, F. S. 1953. The Medusae of the British Isles : Anthomedusae, Leptomedusae, Limno- 

medusae, Trachymedusae and Narcomedusae. Cambridge : pp xii + 530, text-figs. 1-3 19 

and Pis. I-XXXV. 
Sars, G. O. 1873. Bidrag til Kundskaben om Norges Hydroider. For h. Vidensk.-Selsk. 

Krist. 1872 : 91-150, Tab. II-V. 
Sars, M. 1846. Fauna Littoralis Norvegiae, Heft 1 : 1-94. Christiania. 

1857. Bidrag til Kundskaben om Middelhavets Littoral-Fauna, Reisebemaerkninger 

fra Italien. Nyt. Mag. Naturv. 9 : 1 10-164. 

1877. New and little known Coelenterates. Fauna Littoralis Norvegiae, Heft 3 : 1—32. 

Schneider, K. C. 1897. Hydropolypen von Rovigno, nebst Uebersicht iiber das system der 

Hydropolypen im allgemeinen. Zool. Jb. (Syst.) 10 : 472-555. 
Steche, O. 1906. Bemerkungen uber pelagische Hydroiden kolonien. Zool. Anz. 31 : 

Stechow, E. i 91 9. Zur Kenntnis der Hydroidenfauna des Mittelmeeres, Amerikas und 

anderer Gebiete, nebst Angaben iiber einige Kirchenpauer'sche Typen von Plumulariden. 

Zool. Jb. {Syst.) 42 : 1-172. 

1 92 1 . Neue Genera und Species von Hydrozoen und anderen Evertebraten. A rch . Natur- 

gesch. 87. A3 : 248-265. 

1922. Zur Systematik der Hydrozoen, Stromatoporen, Siphonophoren, Anthozoen und 

Ctenophoren. Arch. Naturgesch., 88, A3 : 1 41-155. 

1923. Zur Kenntnis der Hydroidenfauna des Mittelmeeres, Amerikas und anderer 

Gebiete. II. Tiel. Zool. Jb. (Syst.) 47 : 29-270. 

Thornely, L. R. 1908. Reports on the marine biology of the Sudanese Red Sea. X. Hyd- 

roida collected by Mr. C. Crossland from October 1904 to May, 1905. /. Linn. Soc. Zool., 

31 : 80-85. 
Torrey, H. B. 1902. The Hydroida of the Pacific coast of North America. Univ. Calif. 

Publ. Zool. 1 : 1-104. 
Totton, A. K. 1930. Coelenterata. Part V. Hydroida. Brit. Antarct. (Terra Nova) Exped. 

1910, Zool. 5 : 131-252. 
Uchida, T. 1927. Studies on Japanese Hydromedusae. 1. Anthomedusae. /. Fac. Sci. 

Tokyo Univ. 1 : 145-238. 
Vanhoffen, E. 1910. Die Hydroiden der deutschen Sudpolar-Expedition 1901-1903. 

Dtsch. Sudpol. Exped., 11, Zool. 3 : 269-340. 
Vervoort, W. 1946. Hydrozoa (C.i). a. Hydropolypen. Fauna van Nederland, 14: 

Weismann, F. L. A. 1883. Die Entstehung der Sexualzellen bei den Hydromedusen. Zugleich 

ein Beitrag zur Kenntnis des Baues und der Lebenserscheinungen dieser Gruppe. Jena : 

pp. xiii -f 295, 24 pis. 
Wright, T. S. 1857. Observations on British Zoophytes. Edin. New Phil. J., N.s., 6 : 


1858. Observations on British Zoophytes. Proc. R. phys. Soc. Edinb., 1 : 447-455. 

1 861. Observations on British Protozoa and Zoophytes. Ann. Mag. Nat. Hist. (3) 

8 : 120-135. 

1867. Observations on British Zoophytes . . . Proc. R. phys. Soc. Edinb. 3 : 42-46. 



(With Plates 11-12) 


Three species of Hydractinia, H. allmani Bonnevie, H. carica Bergh and H. monocarpa Allman 

from the Arctic are redescribed from numerous colonies. The great differences noted between 

colonies of the same species are shown to depend on the stage of reproductive exhaustion reached 

when the colonies were taken. What is known of their biology and distribution is summarized. 


Hydractinia is a large genus, and, of the many species which have been described, 
few have been worked on alive prior to a description being drawn up. In consequence 
many of the specialists who have worked on these animals (perforce from Expedition 
material) have seldom considered the specimens before them as having been something 
alive and constantly changing. Inevitably phases in growth and reproduction as well 
as ecological forms have on occasion been described as distinct species. 

The northern Hydractinia species have been partially revised by Broch (1916) and 
Kramp (1932) so that there is now less confusion about the identity of arctic and 
subarctic forms. Some species however require redescription based on re-examination 
of as many colonies as possible in order to eliminate the use of so-called diagnostic 
features which are due to growth, seasonal variation, reproductive activities or 
reflect environmental conditions. 

In species like Coryne where the gonophores are borne directly on the body of the 
hydranth, these reproductive activities cause the hydranth itself to be resorbed and 
often reduced to a mere stump without tentacles. This phenomenon may be termed 
reproductive exhaustion and it is by no means confined to the capitate hydroids 
where its effects are most noticeable. In Hydractinia reproductive exhaustion mani- 
fests itself first in the fertile polyps (or special reproductive polyps as the case may be) 
and also causes some reduction in the size of the nutritive polyps. I first noticed an 
appreciable reduction in the size of the sterile polyps in a Hydractinid due to this 
cause in the undescribed hydroid of Podocoryne borealis. During the fifteen days I had 
it under observation, it was producing large numbers of medusae, and by the time 
I was certain that it was distinct from the better-known P. carnea, the colony had 
altered so much in appearance that I could not honestly describe it in that condition 
(Rees, 1941). 


This paper is confined to observations on three northern species, Hydractinia 
allmani Bonnevie, Hydractinia carica Bergh and Hydractinia monocarpa Allman, and 
is based on the rich arctic and subarctic collections of the Naturhistoriska Riksmuseum, 
Stockholm. Some of the material has been briefly described by Elof Jaderholm, but 
all the locality records of colonies in the Riksmuseum have been reported by him in 
various papers. 

Hydractinia allmani Bonnevie 

Hydractinia allmanii Bonnevie, 1898, Z. wiss. Zool., 63 : 485, pi. xxvi, figs. 36-37. 

Bonnevie, 1899, Norske Nordhavs Exped., 1876-8, 26 : 47, pi. i, fig. 1. 

Jaderholm, 1902, Bih. Svenska Vet. Akad. Handl. 28, Afd. 4, No. 12 : 8. 

Broch, 1903, Bergens Mus. Arb. 1903, No. 9, Table. 
Hydractinia allmani, Hartlaub, 1905, Zool. Jb. Suppl. 6, Bd. 3 : 518. 

Jaderholm, 1908, Mem. Acad. Sci. St. Petersb. (8) 18, No. 12 : 7, pi. i, fig. 5 ; pi. ii, figs. 


Jaderholm, 1909, K. Svenska Vetensk-Akad. Handl. 45, No. 1 : 49. 

Broch, 1910, Fauna Arctica, 5 : 200. 

Kiihn, 1910, Zool. Jb. (Anat.), 30 : 107. 

Kramp, 1914, Medd. Gronland, 23 : 988. 

Broch, 1916, Danish Ingolf Exped. 5 : 44. 

Rylov, 1923, Annu. Mus. Zool. Acad. St. Petersb., 24 : 151. 

Kramp, 1943, Medd. Gronland, 121, No. 11:8. 

Type locality. Norske Nordhavs Expedition 1876-78, St. 137, 67 24' N., 
8° 58' E., 827 m. (Zoological Museum, Oslo) Type locality selected here. 

Material seen. In Riksmuseum, Stockholm : 

Swedish Polar Expedition, 1900. 

St. 17, Mackenzie Bay, north of Franz Josef Fjord, E. Greenland, 12-35 m - 

St. 23, Mackenzie Bay, north of Franz Josef Fjord, E. Greenland, 3-10 m. 

St. 27, Muskoxfjord, Franz Josef Fjord, E. Greenland, 220 m. 
Russian Polar Expedition, 1 900-1 903. 

St. 10 d, Kara Sea, 73 27' N., 79 15' E., 40 m. 

St. 18, Gulf of Middendorff, 75 54' N., 92 59' E., 12-14 m. 

St. 50, north of New Siberian Isles, 77 20' 30" N., 138 47' E., 38 m. 
Otto Torell : Bellsund, Spitzbergen, 8-12 fm. 

Specific Characters. Hydractinia with encrusting base, with few simple, smooth 
spines. No tentaculozooids or spiral zooids. Nutritive polyps, long columnar 
(1-4-15-0 mm. high) with up to 16 filiform tentacles. Reproductive polyps (075-5-0 
mm.) at first with hypostome and at least 8 tentacles becoming reduced to stumps 
with armed proboscis and even without tentacles. Usually one large and one small 
gonophore on each reproductive polyp. 

Gonophores (o-6 — 1-4 X 0-5 — 1-3 mm.). Female, cryptomedusoid with 4 well- 
developed radial canals and 8 teat-like tentacle rudiments. Eggs numerous, polygonal 
at first becoming rounded or oval and escaping through a pore between the tentacle 
rudiments. Male gonophores also cryptomedusoid. 


Distribution. Apart from two records in rather deep water off the Norwegian 
coast, viz : position, 67 24' N., 8° 58' W., S.W. of the Lofotens at a depth of 827 m. 
(Bonnevie, 1899) and position 63 12' N., i° 30' E., at a depth of 1320 m. (Broch, 
1903), H. allmani has been found mainly in the high Arctic. 

It has been found in the Spitzbergen area on a number of occasions (Bonnevie, 
1899 ; Jaderholm, 1909 and Rylov, 1923). To the east the Russian Polar Expedition 
1900-1903 took it in the Kara Sea, the Gulf of Middendorff and north of the New 
Siberian Islands (Jaderholm, 1908). In Greenland it was taken at three stations by 
the Swedish Polar Expedition, 1900, in the neighbourhood of Franz Josef Fjord 
(Jaderholm, 1902) and in Kempes Fjord by the Swedish Gronland Expedition, 1899. 
Other Greenland records from Hurry Fjord (Scoresby Sound area), Kempes Fjord, 
Vega Sound and the mouth of Forsblad Fjord are given by Kramp (1943). 

The absence of records from the Okhotsk Sea and the Bering Sea are possibly due 
to lack of observations and the species may prove to be circumpolar in distribution. 

Description. This is a large and handsome Hydractinia, but small and intermediate 
colonies occur and the species is very variable. For this reason several colonies are 
described below. 

Fully grown colonies before the onset of reproductive exhaustion, as already 
mentioned above, are large and handsome and the following description is based on a 
colony from Muskoxfjord, East Greenland (PL 1 1 , figs. 3-4). The encrusting base, which 
is continuous, has few spines, and arising from it are long spindly nutritive polyps 
reaching a height (to hypostome) of 8-0-13-5 mm. in preserved material. There is a 
well developed, conical proboscis, which is armed with a few nematocysts, with at 
its base, a circle of about 15 tentacles in a close double whorl. Each hydranth arises 
directly from the hydrorhiza and there is no distinct ring of perisarc at this point 
although the perisarc forms a thin wrinkled sheath. 

The largest reproductive hydranths are 1-6-4-5 mm. high, that is, they are only 
about half the length of the nutritive ones, and from this colony it is not possible to 
say whether they are derived from nutritive polyps. The largest seen had a length of 
4-5 mm.; it had a conical proboscis moderately well armed with nematocysts at its 
distal end. The tentacles were four in number and only 0-2 mm. in length. Some 
distance below the head of the hydranth it carried one large and one small gonophore. 
Other blastostyles ranged down to i-6 mm. in height and the tentacles were some- 
times reduced to mere knobs. Each blast ostyle usually carried one large and one 
rudimentary gonophore. None of the gonophores was fully developed, the largest 
having a diameter of 1-4 mm. No spiral zooids have been observed. 

Both male and female gonophores occur in the same colony. They are cryptomedu- 
soids with well developed radial canals and the tentacle bulb rudiments are repre- 
sented by four slightly pigmented areas (more mature gonophores are described from 
the colony taken at Mackenzie Bay, East Greenland on 1st August, 1900). 

Another colony from Mackenzie Bay (also East Greenland) has similar gonophores, 
but the nutritive polyps are either less well developed or retracted (due to preserva- 
tion methods) (PL 1 1 , figs. 1 and 2). The polyps have a height of up to 5 mm. Likewise 
the gonophores are not fully mature and the blastostyles themselves are quite small 
being only 1*3-1-5 mm. in total height. 



A second colony from Mackenzie Bay taken on ist August, 1900, has distinctly 
smaller hydranths, these being less than 3-0 mm. in height, but both male and female 
gonophores are fully developed. The female gonophore is fully developed at 1-2 mm. 
in diameter and the tentacular rudiments are represented by 4 - 8 teat-like processes 
and the manubrium may protrude through the opening by which the eggs escape. 

Another colony taken by the Russian Polar Expedition, 1900-1903, St. 50, north 
of the New Siberian Islands, is remarkable for the length of the nutritive polyps 
which are up to 14-3 mm. in height. The female gonophores are ripe (PL 11, fig. 7) and 
the male ones nearly so. The blast ostyles vary from 1-5 mm. to 3-3 mm. in height 
and the larger carry 8 tentacles and the smaller ones only 1 or 2. Typically only 2 
gonophores, one large and one small, are found on each blastostyle. The ripe female 
gonophore has 4 perradial rudiments of tentacles and 4 smaller interradial rudiments 

(pi. 11, fig. 7 ). 

The individual (both nutritive and reproductive) polyps of a colony from the Kara 
Sea are small. Some of the female gonophores are sufficiently advanced to show the 
rudiments of tentacle bulbs and radial canals. Here the blastostyles are more reduced 
than in some of the other colonies and often only one or two stumps of tentacles are 
left (PI. 11, figs. 5 and 6). The proboscis of the reduced blastostyle is well armed with 
nematocysts. Eggs are numerous at first but become reduced in number to 10-15 
in the ripe gonophore. 

As will be noticed from Table I there is a considerable range in the size of the 
nutritive polyps and the very large size of the polyps in two colonies (one from 
Mackenzie Bay, St. 27 and the other from north of the New Siberian Islands) made 
me suspect that a different species was involved, but a careful re-examination of the 
material revealed no essential points of difference in any of the colonies except in size 
and degree of maturity of the gonophores. 

Table I. — Hyd? -actinia allmani Bonne vie. 
(Measurements in mm.) 

Length of nutritive polyp 
Maximum diameter of polyp 
Length of hypostome 
Height of reproductive polyp 
Length of gonophore 
Diameter of gonophore . 

Mackenzie Bay, E. 


Gulf of 

North of 



New Siberian 



St. 17 

St. 23 

St. 27 

Kara Sea 


. 2-4-2-7 

1 -4-3- 1 


2 • 0-2 • 2 


. 0-4-0-5 

• 2-0 • 4 


• 2-0 • 7 

• 7-0 • 9 

. 0-5-0-6 


• 6-0 • 8 


• 5-0 • 8 

• 1 -5-i -7 

1 -1-2-9 




. 1-1-1-3 

• 9-1 • I 

o-6-i -4 

• 6-1 • 1 

1 -i-i -4 

. I-Q-I-2 

• 9-1 • I 




In these colonies the nutritive polyps are more than twice the size of the largest 
reproductive polyps ; the latter have up to 8 tentacles and are very similar to the 
sterile polyps. It is not possible to be completely certain that they are reduced from 
the nutritive polyps through bearing gonophores, as in Podocoryne carnea, but it may 
be that we have here the beginnings of the specialization which results in the pro- 
duction of the specialized reduced reproductive polyps seen in Hydractinia echinata. 


In the remaining colonies there is some overlap in the size of the two kinds of polyps, 
and in view of the small size of all the polyps it appears likely that they are reduced 
by prolonged breeding. This raises the question of the status of Hydr actinia ornata 
Bonne vie, which, as far as I can see, is distinguished from H. allmani only on size ; 
its status must await the re-investigation of the original material. 

There is one feature in the reduction of the reproductive polyp which is worth 
noting. The proboscis of the very reduced polyp is armed with nematocysts, these 
being concentrated at its tip as the tentacles are resorbed. The use of this character 
(an armed proboscis) in classification should therefore be used with caution. 

Biology. In all records where the substratum of Hydr actinia allmani has been 
indicated, the gastropod involved has been Sipho curtus (Jeffreys). In the high Arctic 
the hydroid occurs at depths of 3-760 m. in Greenland, 75-350 m. off Spitzbergen 
and in 12-40 m. in the more easterly localities. 

Records of this species from off the Norwegian coast in comparatively deep water 
(see Distribution) suggest that this submergence is related to temperature. A colony 
reported by Jaderholm (1909, p. 49) was dredged from 350 m. where the bottom 
temperature was 273 C, while Bonnevie (1899, p. 11) recorded the species from 
bottoms where the temperatures were — i° C. and — 1-2° C. Comparable tempera- 
tures (given by Ekman, 1953, p. 166) exist at the other localities at which this 
species has been found. 

It is probable that Hydractinia allmani can be regarded as a stenothermal cold 
water species. 

Historical and Relationships. Since Bonnevie described the species in 
1898, numerous records indicate that this is a common species in the high Arctic. 
It can be readily distinguished from Hydractinia monocarpa and H. echinata by its 
simple smooth spines ; from Podocoryne it is separated by its fixed gonophores. 
Compared with H. carica, it is more robust in habit, with longer polyps and more 
polyp-like reproductive zooids. The numerous eggs in the female gonophores (eggs 
are few in number in the gonophores of H. carica) also help to distinguish it. 

Hydractinia carica Bergh 1887 

Hydractinia carica Bergh, 1887, Lutken, C.F. Dijmphna-Togtets zool.-bot. Ubbytte, p. 331, pi. 
xxviii, fig. 1. 
Schydlowsky, 1902, Trav. Soc. Nat. Univ. Kharkow, 36 : 114. 
Hartlaub, 1905, Zool. Jb. Suppl., 6, Bd. 3 : 518. 
Jaderholm, 1908, Mem. Acad. Sci. St. Petersb. (8) 18, No. 12:8. 

Jaderholm, 1909, K. Svenska Vetens.-Akad. Handl., 45, No. 1 : p. 48, pi. ii, figs. 10 and 11. 
Broch, 1 910, Fauna Arctica, 5 : 200. 

Broch, 1916, Danish I ngolf Expedition, 5, Part 6 : 48, pi. i, fig. 11. 
Scheuring, 1922, Wiss. Meeresuntersuch. Abt. Helgoland, 13 : p. 168. 
Rylov, 1923, Annu. Mus. Zool. Acad. St. Peterb., 24 : 150, Taf. vi, figs. 2-3. 
Fraser, 1931, Contr. Canad. Biol., n.s., 6 : 6. 
Uschakow, 1937, Trans. A ret. Inst. Leningr. 50 : 12. 
Fraser, 1944, Hydroids Atlant. Coast N. America, p. 77, pi. xiii, fig. 53. 


Hydractinia minuta Bonnevie, 1898, Z. wiss. Z00L, 63 : 468, pi. xxvi, fig. 38. 
Bonnevie, 1899, Norske Nordhavs Exped. 1876-8, 26, Zool. : 47 pi. i, fig. 3. 
Bonnevie, 1901, Meeresfauna von Bergen. Hydv., 1 : 7. 
Broch, 1910, Fauna Arctica, 5 : p. 141 and 200. 

Type locality. Petuchoffskoi Schar, Kara Sea, 15 m. (Dijmphna Expedition — 
Zoological Museum, Copenhagen) . 

Material seen. In Riksmuseum, Stockholm : 

Fragment of type material on a mounted slide. 

Bellsund, Spitzbergen, 20 fm. 22nd July, 1864 (A. J. Malmgren) 

Bellsund, Spitzbergen, 8-12 fm. (Otto Torrell) 

Recherche Bay, Spitzbergen, 13th July, 1898 (Spitzbergen Expedition, 1898) 

67 56' N., 66° 18' W., 13th October, 1897 (E. Nilsson) 
Russian Polar Expedition, 1900-1903, St. 3, Murman Sea, 69 35' N., 50 5' 

E.; 70 m. 

In British Museum : 

Klaas Billen Bay, Spitzbergen, 28th July, 1921, C. S. Elton (Oxford Univ. 
Exped. to Spitzbergen, 1921). 

Specific characters. Hydractinia with encrusting base with few high conical 
smooth spines (up to 07 mm. high) with rounded apices. Nutritive polyps, small, up 
to 2-3 mm. high when fixed, with a single whorl of 10-16 tentacles. Proboscis simple, 
almost unarmed. No spiral zooids known. 

Reproductive polyps up to 2-3 mm. high, at first with conical hypostome and at 
least 8 tentacles, becoming greatly reduced to very short stumps without tentacles ; 
proboscis armed with nematocysts when present. Gonophores 3-7 in number borne 
in a ring round the polyp. 

Gonophores crypt omedusoid, with 4 radial canals and rudiments of tentacles 
present but not developed into bulbs. Eggs large, 4-10 in number. 

Distribution. Hydractinia carica is known from the Kara Sea (Bergh, 1887), 
the Barents Sea (Jaderholm, 1908 ; Scheuring, 1922), Spitzbergen, the Davis Strait 
area (West Greenland) and on the Norwegian coast in the neighbourhood of Bergen. 

There are numerous records of this species from Spitzbergen, most of these being 
summarized by Jaderholm (1909) and Rylov (1923). I have been able to verify 
Jaderholm's record (1908) from Davis Strait and there are others by Fraser (1931 and 
1944 as carica and minuta) from approximately the same area. The most southerly 
record is that of Bonnevie (1901) from the Hjeltefjord, Bergen as H. minuta. 

There is insufficient information to hazard any opinion as to whether this species 
has a circumpolar distribution. 

Description. This is an encrusting species with large spines but without small 
prickles. The large spines are simple, smooth and not sharply pointed ; they have no 
pore at the tip and in size range up to 07 mm. in length. 

In preserved material the hydranths are tubular to fusiform in shape and may be 
over 2-o mm. in height. The head of the hydranth is sometimes clavate and there is a 



conical proboscis without many nematocysts around the mouth. There are usually 
about 16 tentacles arranged more or less in a single whorl. 

The reproductive polyps vary considerably from colony to colony according to the 
stage they had reached when taken, and it has been deemed desirable to describe 
them in the different colonies. 

In the colony from Bellsund, Spitzbergen the reproductive polyps are completely 
reduced to very short stumps 0-17 mm. or less in length, bearing up to 5 gonophores 
each. These have diameters of about 0-45 mm. Sterile hydranths are 1-3-2-0 mm. in 

Another colony from Recherche Bay, Spitzbergen, does not appear to have reached 
the same state of reproductive exhaustion as the one from Bellsund. In this colony 
the fertile polyps are still recognizable as polyps ; one of these still possessed 8 fully 
developed tentacles and a distinct proboscis and carried 5 moderately developed 
gonophores. Another reproductive polyp, with a height of 0-9 mm., had 7-8 tentacles 
and 4 gonophores each with a diameter of 0-3 mm. 

Text-fig. 1. — Hydractinia carica Bergh : blastostyles from a colony from Recherche 
Bay, Spitzbergen (Swedish Spitzbergen Expedition, 1898). 

A colony from position 67 56' N., 66° 18' W., had particularly large blastostyles, 
1-0-2-3 mm - i n height to hypostome. These had 3-5 gonophores each and only a few 
had up to 2 reduced tentacles. The hypostome was usually well armed with nemato- 

There were only a few very reduced polyps. These were so short that they could 
not be measured accurately ; they carried 5-7 gonophores and were clearly reaching 
the exhaustion stage. 

Colonies from the Murmansk coast and Bellsund, Spitzbergen, were fertile and some 
of the reproductive polyps of these possessed 2 or more tentacles. 

A mounted slide marked Orig. Ex., Dijmphna Exp. presented by Levinsen appears 
to consist of part of the type material of this species. Measurements of this and other 
colonies in the Riksmuseum are given in Table II. Some of the reproductive polyps 
carry about 4 tentacles and others have lost them (Text-fig. 1). 





— Hydractinia carica Bergh 

(Measurements in mm.) 

Kara Sea 

(fragment of 

67 56' N., 

Bell Sound, 

type material) 

66° 18' W. 


Length of polyp 

1-5-1 -7 

1 • 0-2 • 3 

1 • 0-2 • 

Maximum diameter of polyp 


• 1-0 • 2 

• 2-0 • 3 

Length of hypostome 

o- 4 


• 1-0 • 3 

Height of reproductive polyp . 

o • 75-0 • 9 


■ 1 7-0 • 2 

Length of gonophore 

o • 28-0 • 4 



Diameter of gonophore 

• 25-0 • 4 



Length of spine 




Only female gonophores have been noted in this material ; they are cryptomedu- 
soids without radial canals but with apical patches of nematocysts. There is a well- 
developed manubrium and when ripe the gonophores contain 4-10 large eggs each 
(Text-fig. 2, a, b). 

Text-fig. 2. — Hydractinia carica Bergh ; gonophores from a fragment of the type colony, 
Kara Sea (Dijmphna Expedition) : a, female gonophore ; b, another female gonophore 
in optical section. 

This material agrees with the earlier descriptions of Bergh (1887), Jaderholm 
(1908) and Broch (1916), but it has been possible to get a better picture of the species 
as a whole. The nutritive polyps (in preserved material) are seldom more than 2-0 
mm. in height and it is possible that the reproductive polyps are derived directly 
from them. Bergh and the other authors indicated that the reproductive polyps 
might have 1-4 tentacles, but I have found 8 which supports my suggestion. Jader- 
holm (1909, pi. ii, figs. 10 and n) gives good illustrations of partially reduced hydranths, 
and others are figured here (Fig. 1), but they may become so short that even the 
hypostome is almost completely reduced and the stalk below the point of origin of 
the gonophores also becomes extremely short. Broch (1 916, p. 48) is undoubtedly 
right in his contention that Bonnevie's H. minuta (Bonnevie, 1899, p. 48) is identical 
with H. carica. I regard the colony she described as being in an advanced stage of 
reproductive exhaustion. 

Biology. This species favours the shells of various northern species of Buccinum 
as a substratum. In the Barents Sea it has been found on Buccinum undatum and 
B. tenue. In the Spitzbergen area Jaderholm (1909) has found it on Buccinum 
glaciate and Trophon clathrus, while Rylov (1923) reported it as H. minuta on Buccinum 
ovum and B. ciliatum. 


The bathymetric range appears to be 0-120 m. in the Spitzbergen area and 40-70 
m. in the Barents Sea. 

Hydractinia carica may be regarded as mainly a cold water species which, if 
Bonnevie's record from Bergen is authentic, is probably less stenothermal than H. 

Historical and Relationships. Hydractinia carica was described by Bergh 
(1887) from Petuchoffskoi Schar in the Kara Sea from a depth of 15 metres. 
Subsequently it was recorded many times in the far north and as H. minuta Bonne vie 
from Spitzbergen and elsewhere. As a species it is distinguished from other northern 
Hydractinia by its simple smooth spines, the extreme reduction which takes place 
in its reproductive polyps and by its gonophores. These have no radial canals and 
the female ones have few eggs. 

Hydractinia monocarpa Allman 

Hydractinia monocarpa Allman, 1874, Nature, Lond. 11, No. 270 : 179. 
Marktanner, 1895, Zool. Jb. (Syst.), 8 : 394. 
Schydlowsky, 1902, Trav. Soc. nat. Univ. Kharkow, 36 : 114. 
Hartlaub, 1905, Zool. Jb. Suppl. 6, Bd. 3 : 518. ' 
Jaderholm, 1908, Mem. Acad. Sci. St. Petersb. (8), CI. Phys.-Math. 18, No. 12 : 8, Taf. 1, 

fig. 6, Taf. 2, figs. 6-9. 
Jaderholm, 1909, K. Svenska Vetens.-Akad. Handl. 45, No. 1 : 49, Taf. ii, figs. 12-13. 
Broch, 1910, Fauna Arctica, 5 : 141. 
Broch, 1916, Danish Ingolf Exped. 5, Pt. 6 : 47. 

Rylov, 1923, Annu. Mus. Zool. Acad. St. Petersb. 24 : 148, Taf. VI, fig. 7. 
Kramp, 1932, Medd. Gronland, 79, No. 1 : 16. 
Uschakow, 1937, Trans. Arct. Inst. Leningr., 50 : 12. 
Fraser, 1944, Hydroids Atlantic Coast N. America, p. 77. 

Hydractinia echinata, Wintrier, 1880, Naturh. Tidsskr. 12 : 227, 254. 

Levinsen, 1893, Vidensk. Medd. naturh. Foren Kobenhavn, 1892, p. 153. 

Type locality. Spitzbergen, on Trophon clathratus L. (Allman, 1876) (Zoological 
Museum, Copenhagen) 

Material seen. In Riksmuseum, Stockholm : 

Matotschkim Schar, 15 fm. (Novaya Zemlya Expedition, 1875). 
Russian Polar Expedition, 1900-1903, St. 53, 77 10' N., 142 48' E. (N. of New 
Siberian Isles) 35 m., on Trophon clathratus and Beta plicij era. 

Specific characters. Hydractinia with encrusting base with long, often irregular, 
hollow spines which may bifurcate distally. Spines longitudinally keeled especially 
at base, but smooth and not thorned, up to 2-4 mm. high. Nutritive polyps fusiform 
to tubular, up to 3-6 mm. high, borne on the encrusting base or on the sides of the 
spines. Tentacles, 10-15. Spiral zooids not known. 

Reproductive polyps, up to 1-5 mm. high, with armed proboscis and at least 4 
tentacles, or may be reduced without tentacles and greatly shortened. Gonophores 
1-4 on each, but usually only one. 


Gonophores : female, up to 1-4 mm. in diameter ; cryptomedusoid with 4 radial 
canals and apical patch of tissue. Male gonophores similar, up to 0-55-0-6 mm. in 

Distribution. All records of this species are from the high Arctic, chiefly from 
Spitzbergen, Novaya Zemlya and the New Siberian Isles. It appears to be a common 
species on shells at Spitzbergen (Airman, 1876 ; Jaderholm, 1909 and 1916 ; and 
Broch, 1910). The Russian Polar Expedition 1900-1903 took it at two stations off 
the New Siberian Islands and in Chatanga Bay in Nordenskj old's Sea (Jaderholm, 
1908). Records from Novaya Zemlya are given by Jaderholm (1909). 

West Greenland records from Upernivik and from near Cape Atholl are given by 
Kramp (1932). This distribution suggests that the species is circumpolar. 

Description. This species has an encrusting base, but the most characteristic 
feature is the presence of long, upright irregularly-formed spines rising to a height of 
about 3 mm. (PL 12, figs. 8-1 1). The spines are seldom straight or symmetrical and 
are usually curved towards the tip which tapers to an open hyaline or horn-coloured 
tube. Frequently the spines are double or bifurcated and their surface is seldom 
smooth. The naked coenosarc may cover a good part of the spines. 

The nutritive hydranths may be long and tubular reaching (when expanded) a 
considerable length. Some preserved examples, moderately well fixed, have a length 
of 3-6 mm., but, when contracted, may only be one- third or less than this length. 
Such retracted polyps are either cylindrical or flask-shaped (PL 12, figs. 9 and 10). 
The hypostome carries nematocysts, particularly near the tip, but they are few in 
number. There are 10-14 tentacles. A curious feature of this species is that one or 
more polyps may be carried on the spines themselves and I have seen up to three on 
one spine (PL 12, figs. 9 and 10). 

The reproductive polyps, noted in the material seen, were up to 1-5 mm. in height 
and were provided with up to 4 tentacles. Each had an armed tip to the proboscis 
and carried up to 4 gonophores. Only female gonophores have been noted by me ; 
these are large, 0-5-0-7 mm. in diameter and are cryptomedusoids with manubrium 
and radial canals. It is clear that the blast ostyles become considerably reduced due 
to reproductive exhaustion. Frequently they are reduced so much that the gonophoi cs 
appears to be almost sessile on the coenosarc. When this happens the upper part cf 
the polyp seems to become larger and more polypoid, but the stalk below the point 
of origin of the gonophore is extremely short. 

In his original description Allman (1874) described the reproductive polyps as 
devoid of tentacles, but tentacles have been noted by Jaderholm (1908). Rylov 
(1923), Kramp (1932) and by myself. The diagnosis of the species (p. 25) differs 
greatly from that of Allman because more colonies have been available for study. 

Biology. Hydradinia monocarpa has been found on the shells of gastropods such as 
Trophon clathrus, Beta plicifera and Buccinum hydrophanum (Jaderholm, 1908, 1916, 
and Kramp, 1932). The hydroid may grow all over the shell as in a specimen of 
Beta plicifera from Kol Bay, Isfjord, Spitzbergen. The nutritive and reproductive 
polyps flourish best in and near the sutures between whorls, but they also occur on 
exposed surfaces and are absent only on abraded surfaces of the shell. The tall spines 
are fairly closely grouped and must offer some protection to the zooids. 


The species has a known bathymetric range of 14-165 metres, the deepest record 
being from West Greenland (Kramp, 1932). Most of the records from Spitzbergen 
and elsewhere in the Polar Sea are, however, from shallow coastal waters. 

Hydractinia monocarpa may be regarded as a high Arctic or stenothermal cold 
water species. 

Historical and Relationships. Hydractinia monocarpa was described by 
Allman from a colony labelled " Spitzbergen " in the Zoologiske Museum, Copen- 
hagen. It has since been found frequently in the Spitzbergen area and elsewhere in 
the high Arctic. 

This distinctive species, with its long, hollow, often bifurcated spines, its polyps 
arising from both encrusting base and spines, its cryptomedusoid gonophores, with 
four radial canals but without tentacle rudiments, cannot be mistaken for any other 
northern species. 

The presence of hydranths on the spines recalls the much greater development of 
this feature in Hydrissa sodalis (Stimpson) from East Asiatic seas. 


I wish to thank Dr. Bengt Hubendick for hospitality while examining this material 
at the Riksmuseum, Stockholm, and also for the loan of some of it for more detailed 
examination in London. I also wish to thank Mr. M. G. Sawyers for the care he has 
taken in preparing the photographs of Hydractinia allmani and H. monocarpa from 
slides made by the late Elof Jaderholm. 


Colonies of Hydractinia allmani show marked variation according to the stage 
of reproductive exhaustion reached at time of capture and it has been suggested here 
that the reproductive polyps in this species are derived from nutritive ones. The 
great variation in the size of nutritive polyps and in the reproductive polyps (from 
well formed ones to almost blastostyles) is highly significant, and implies that we 
must use these features with great caution in distinguishing species. It brings to mind 
the very large number of boreal and tropical species of Hydractinia which have been 
described on single colonies. A critical study of these must result in a great reduction 
in the number of recognizable species. Another feature, noted in this paper, is the 
concentration of nematocysts in the tip of the hypostome as the tentacles are resorbed 
during reduction of the polyp. It means that the armed proboscis, too, must be used 
with discretion as a diagnostic character. 

The study of Hydractinia carica also bears out the general conclusions noted 
above, but this species is notable for the almost complete resorption of the repro- 
ductive polyp and shortening of its stem, so that the gonophores sometimes give the 
appearance of being sessile on the hydrorhiza and even the hypostome may disappear. 
This, too, is a factor to be considered in classification. 

Although H. monocarpa can be readily distinguished by its spines, its reproductive 
polyps vary according to phase and bear out the general conclusions reached for the 
other two species. 


Allman, G. J. 1874. On the diagnosis of new genera and species of hydroids. Nature, 
Lond. 11, No. 270 : 179. 

1876. Diagnosis of new genera and species of Hydroida. /. Linn. Soc. Zool. 10 : 251- 

284, pi. ix-xxiii. 

Bergh, R. S. 1887. Goplepolyper (Hydroider fra Kara-Havet) Liitken, C.F. Dijmphna- 

Togtets. zool.-bot. Udbytte, pp. 329-338, pi. 28. 
Bonnevie, K. 1898. Zur Systematik der Hydroiden. Z. wiss. Zool. 63 : 465-495, pi. 25-27. 

1899. Hydroida. Den Notske Nordhavs Expedition, 1876-1878, 26 : Zool., 1-104, pi. 1-8. 

1 901. Hydroiden. Bergens Mus. Meeresfauna von Bergen, Heft 1 : 1-15, pl.i. 

Broch, Hj. 1903. Die von dem norwegischen Fischereidampfer " Michael Sars " in den 

Jahren 1 900-1 902 in dem Nordmeer gesammelten Hydroiden. Bergens Mus. Arb. 
1903, No. 9 : 1-14, 4 pis. 1 table. 

1910. Die Hydroiden der Arktischen Meere. Fauna Arctica, 5 : 127-248, pi. 2-4. 

1916. Hydroida (Part 1) Danish Ingolf Exped. 5, Part 6 : 1-66, PI. i-ii and 20 text-rigs. 

Ekman, S. 1953. Zoogeography of the Sea. London : Sidgwick & Jackson, pp. xiv + 417. 
Fraser, C. M. 193 1. Hydroids of Hudson Bay and Hudson Strait. Contr. Canad. Biol., 

n.s. 6 : 477-481. 

1944. Hydroids of the Atlantic Coast of North America. Toronto, pp. 454, 94 pis. 

Hartlaub, C. 1905. Die Hydroiden der magalhaensischen Region und chilenischen Kuste. 

Zool. Jb. Suppl. 6, Bd. 3 : 497-714, 142 text-figs. 
Jaderholm, E. 1902. Die Hydroiden der Schwedischen Zoologischen Polar-expedition 1900. 
Bih. K. Svenska Vet.-Akad. Handl. 28, Afd. 4, No. 12 : 1-11, Taf. 1. 

1908. Die Hydroiden des Sibirischen Eismeeres, gesammelt von der Russischen Polar- 
Expedition 1900-1903. Mem. Acad. Sci. St. Petersb. (8), CI. Phys.-Math. 18, No. 12 : 
1-28, pi. i-iii. 

1909. Northern and Arctic Invertebrates in the collection of the Swedish State Museum 

(Riksmuseum) . IV. Hydroiden. K. Svenska Vetens.-Akad., Handl. 45, No. 1 : 1-124, 
pi. i-xii. 

1916. Die Hydroiden des Eisfjords. K. Svenska Vetens.-Akad. Handl. 54, No. 4 

1-14, 1 text-fig. 

Kramp, P. L. 1914. Hydroider. Conspectus Faunae Groenlandicae. Medd. Gvenland, 
23 : 383-456. 

1932. The Godthaab Expedition 1928. Hydroids. Medd. Gronland, 79, No. 1 : 

1-86, 34 text-figs. 

1943- The Zoology of East Greenland. Hydroida. Medd. Grenland, 121, No. 11 : 

1-52, 4 text-figs. 

Levinsen, G. M. R. 1893. Meduser, Ctenophorer og Hydroider fra Gronlands Vestkyst. 

Vidensk. Medd. naturh. Foren Kbh. (5), 4 : 143-212, pi. 5-8. 
Marktanner-Turneretscher, G. 1895. Hydroiden. Zool. Jb. (Syst.) 8 : 391-438, pi. 

Rees, W. J. 1 94 1. On the life history and developmental stages of the medusa Podocoryne 

borealis. J. Mar. biol. Ass. U.K., 25: 129-141. 
Rylov, V. M. 1923. Zoologische Ergebnisse der Russichen Expeditionen nach Spitzbergen. 

Hydroid-Athecata. Annu. Mus. Zool. Acad. St. Peterb. 24 : 140-160, pi. vi. 
Scheuring, L. 1922. Untersuchungsfahrt des Reichsforschungsdampfers " Poseidon " in 

das Barentsmeer im Juni and Juli 1913. Wiss. Meeresuntersuch Abt. Helgoland, N.F. 13 : 

159-183, Taf. V and text-figs. A-C. 
Schydlowsky, A. 1902. Materiaux relatifs a la faune des polypes Hydraires des mers arc- 

tiques. I. Les Hydraires de la Mer Blanche le long du littoral des lies Solowetsky. Trav. 

Soc. Nat. Univ. Kharkow, 36 : 1-276, Taf. I-V. 
Uschakow, P. 1937. Materials on the Hydroids of the Arctic Seas of the U.S.S.R. Trans. 

Arct. Inst. Leningr. 50 : 5-34, 4 text-figs. 
Winther, G. 1880. Fortegnelse over de i Denmark og dets nordlige Bilande fundne Hydroide. 

Naturh. Tidsskr. 12 : 223-278. 


Hydractinia allmani Bonne vie 

Figs, i and 2. — Fragments of a colony from St. 23, Mackenzie Bay, East Greenland (Swedish 
Zoological Polar Expedition, 1900) ; x 10. 

Figs. 3 and 4. — Fragments of a colony from St. 27, Muskoxfjord, Franz Josef Fjord (Swedish 
Zoological Polar Expedition, 1900) ; x 5. 

Figs. 5 and 6. — Reproductive zooids with female gonophores ; St. 18, Karahavet, Midden- 
dorff's Sea (Russian Polar Expedition, 1 900-1 903) ; X 25. 

Fig. 7. — Reproductive zooid with ripe female gonophore ; St. 50, north of New Siberian 
Islands (Russian Polar Expedition, 1900-1903) ; x 14. 

Bull. B.M. (N.H.) Zool. 3, 8. 

PLATE 1 1 , 

Hvdr actinia allmani Born 

Hydr actinia monocavpa Allman from St. 53, north of New Siberian Islands (Russian Polar 
Expedition, 1900-1903). 

Fig. 8. — Spines and female gonophores ; X 12. 

Figs. 9-11. — Spines and nutritive polyps; x 20, x 21, X 25. 

Bull. 11. M. (A\M.) Zool. 3, 8. 




Hydractinia monocarpa Allman, 

r 2 MAR 1956 


2 MAR 1956 





ZOOLOGY Vol. 3 No. 9 

LONDON: 1956 






Pp- 363-385 ; 7 Text-figures 

ZOOLOGY Vol. 3 No. 9 

LONDON: 1956 

(NATURAL HISTORY), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical Series. 

Parts will appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

This paper is Vol. 3, No. 9 of the Zoological series. 


Issued February, 1956 Price Six Shillings 





The genus Ophioseps cannot be maintained ; its past recognition was due to errors of identi- 
fication and observation. Aprasia is the most completely adapted of the Pygopodidae for 
fossorial life, the family exhibiting a continuous series in degeneration of limbs and girdles, with 
the development of a serpentiform habitus and cranial modifications for burrowing. There 
are three species with a combined range from Victoria to the Monte Bello Islands but which 
occur together in only a restricted area near Albany. Two of the species show a considerable 
range of variation in the numbers of presacral vertebrae and scales. This variation appears 
to be correlated with differing summer temperatures. Since the characters are meristic and 
such characters have been shown, in other cold-blooded animals, to be directly affected by the 
temperature prevailing during embryological development, it is suggested that the variation 
may prove to be thermoplastic. The other taxonomically useful, but non-meristic, characters 
show no significant variation in different climatic regions. The temperature correlated clines 
are discontinuous geographically, permitting recognition of two new subspecies. 

Two small limbless lizards were collected on Hermite Island, one of the Monte 
Bello group, in 1952. They differ from any of the known species of the genus Aprasia 
but most nearly resemble Aprasia repens Fry. Since the Monte Bello Islands are 
geographically far removed from the recorded range of any species of Aprasia a 
re-survey of the known species of the genus, and its alleged ally Ophioseps, was 
necessary to ascertain the probable significance of the differences. More than 120 
specimens were assembled for comparison, and this series covered not only the 
whole of the previously known geographical range, but included recently captured 
specimens from regions where none of these lizards had previously been reported. 


As will be demonstrated presently the series is subdivisible into five groups with 
morphological differences, but all the specimens share the following characters : 

Scales smooth, subequal dorsally and ventrally except on the throat and pectoral 
region, where they are slightly smaller than on the belly ; 12 or 14 longitudinal 
rows of scales at mid-body. General habitus worm-like, the head and tail not being 
wider or narrower than the body, which is very elongate and cylindrical. The 
snout projects in front of the labial margin and the large rostral shield is visible 

ZOOL. 3, 9. 24 



from above ; five supralabials, the first being wholly or partly fused with a large 
nasal shield, which forms a suture with its fellow behind the rostral ; pre-frontals 
large, forming a median suture and in contact with the second labial ; a small 
pre-ocular in contact with the third labial and the supra-ocular, which reaches the 
fourth supra-labial or is separated from it by a single small post-ocular ; frontal 
hexagonal ; parietals not much larger than the scales of the occiput and nape, not 
in contact behind the frontal. Eye covered by a circular brill edged with a ring 
of small granules. Shields and scales of the head beset with numerous minute 
pustules, presumably tactile sense-organs. No external auditory meatus, tympanum 
or tympanic cavity. No vestiges of fore-limbs externally ; hind-limbs visible as 
small triangular flaps at the lateral corners of the vent, each being covered by a 
single triangular scale not very different in size or appearance from the other scales 
of the region. No pre-anal pores. A pair of post-anal sacs close behind the vent 
and opening by a pair of small pores just within its posterior border. Bordering the 
anterior margin of each of these sacs there is, in males, a post-cloacal bone which 
pierces the skin laterally close, but caudomesial, to the hind-limb vestige ; the 
protruding portion is sometimes claw-shaped, with the free terminal part of the claw 
directed postero-dorsally, or sometimes subcircular in surface view (Text-fig. i). 

The skull of Aprasia repens has recently been figured by McDowell & Bogert 
(1954, fig. 21) and specimens of all the species represented in the present material 
(vide infra) have been found to conform to this pattern. There is a single pre- 

Fig. 1. Pelvic region of a cleared specimen seen in three-quarters ventro-lateral view 
from the right side to show hind limb, girdle and post-cloacal bones. (Aprasia striolata, 


maxilla ; paired nasals form a long median suture ; the prefrontals are separated 
mesially and extend posteriorly above the orbit to form a suture with a small post- 
frontal ; the frontals are fused to form a single median element, but the parietals 
are paired, forming a long median suture ; there is a narrow supra temporal 
(tabular). 1 In both sexes the maxillary is edentulous and the dentary is toothed ; 
the premaxilla is toothed in adult males of all species, but is completely edentulous 
in juveniles of all species and also in females, except in A. striolata where vestigial 
teeth may be present. In the lower jaw the splenial is wanting or is fused with the 
prearticular. The only skull examined that deviates in any way from this pattern 
is the one figured and described by Jensen (1901) as Ophiopsiseps nasutus Bocage. 
This description has been the cause of much confusion. The lizard was incorrectly 
identified and the description is incorrect in many points that subsequent authors 
have accepted as indicating the existence of a genus (Ophioseps or Ophiopsiseps) 
distinct from Aprasia. As will be demonstrated presently (p. 380) , the type specimens 
of the type species of Aprasia Gray and Ophioseps Bocage are conspecific and the 
two names are thus strict synonyms. But even apart from this nomenclatorial 
consideration there appear to be no grounds for the recognition of a second genus. 
Jensen's specimens have been compared with a series of others from the same 
geographical area (Victoria and S. Australia) and have been found to be juveniles 
of the one and only species that occurs there. This, in its adult phase, has a skull 
like that described above. Jensen was in error in several respects, notably in 
regard to the frontals, prefrontals, postfrontals and " squamosal " (supratemporal) 
which, despite the immaturity of the skull, do not differ from those of the adult. 
He was also misled by the absence of any median suture between the parietals 
into believing that these bones were completely fused ; in fact, owing to immaturity, 
the development of the parietals is incomplete, but they exist as two very thin 
lateral blades widely separated on the mid-line. 

Although all the species and subspecies examined have skulls that are essentially 
similar in their general structure, there will undoubtedly be found to be specific 
differences and, perhaps, ontogenetic differences in proportion. Insufficient skeletal 
material is available to the writer to permit of any definite conclusions, but in the 
long-snouted species (A. repens) the pre-orbital increase in length is accompanied 
by an increase in length of the nasals. Thus, the ratio of frontal to nasal length 
was found to be approximately 1-24 (c.f. also Bogert & McDowell's figure where the 
ratio is 1-25) in this species, but 1-40-1-49 in specimens from South Australia 
{A. striolata) and 1-50-1-56 in an example of Aprasia pulchella and the holotype of 
Ophioseps nasutus Bocage. 

Other skeletal features of the series are as follows. Presacral vertebrae 88-137, 
the number varying from species to species ; the first three vertebrae (four in one 
specimen) lack ribs ; sacral vertebrae three, with sacral ribs diminishing in length 
caudad, the anterior two being free and the last knob-like and fused to the centrum 
(Text-fig. 2). Shoulder girdle a V-shaped structure composed of a pair of rod-like 

1 McDowell & Bogert (1954) figure this bone in their dorsal and ventral views (figs. 21 a and b) but 
do not label it ; it is not shown in the lateral view (fig. 21c). 

zool. 3, 9, 24§ 


coracoids connected across the mid- ventral line by a V-shaped sternal cartilage. 
Each coracoid, which is slightly expanded at its ends, runs antero-dorsally from 
the sternum to a point near the distal extremity of the first rib. The dorsal end 
of each is tipped with a short, tapering length of cartilage which may represent 
a scapula. There are no traces of any bones of the fore-limb. 

The pelvis (Text-figs, i and 2) consists of a subtriangular, slightly curled ilium 
and an ischiopubis. The ilium articulates at its postero-dorsal corner with the 

Fig. 2. Aprasia striolata striolata. Left pelvis and hind limb of a male, diagrammatic 

to show sacral connexions. 

third, knob-like, sacral rib and the other two sacral ribs have loose ligamentous 
connexions with its antero-dorsal border. The ischium and pubis are fused, the 
ischiadic portion of the combined bone being rectangular and the pubic portion 
rod-like ; both portions are widely separated from their fellows on the mid-ventral 
line. The hind limb consists of a short, stout femur, with a well-developed trochanter, 
and a nodule-like vestige of a tibia ; sometimes there is also a vestigial fibula. 


The various forms possessing these characters in common are clearly a closely 
related group and this position is best expressed taxonomically by treating them 
as a single genus. McDowell & Bogert (1954) in discussing the relationships of the 
family Pygopodidae have pointed out that " the more typical pygopods, the Pygo- 
podinae and Lialinae " are akin to the geckoes in many features. But Jensen's 
account of Ophioseps misled them and, although they do not commit themselves 
definitely, they say " we see no reason to question Jensen's placing of Ophioseps 
in a distinct family, as Boulenger (1885) suggested. It is probable, however, that 
Aprasia should be transferred to the Ophiosepidae." Since Ophioseps, sensu 
Jensen, is a fallacy any such disposal of Aprasia would be absurd and it remains 


to consider its position in relation to the other genera of the Pygopodidae. It 
shares with Pygopus, Paradelma, Delma and Lialis a unique character, a pair of 
post-cloacal bones forming external hooks as a secondary sex character of the male. 
These structures were first described and figured in 1870 by Fiirbringer in Lialis 
(burtoni) but without the realization that they were associated with sex ; the same 
author also described the limbs and girdles of Pygopus, but found no such structures 
because, presumably, his specimen was a female. Boulenger (1885 : 239) mentioned, 
as a family character, the presence of a spur laterally behind the vent, but wrongly 
described it as a projection of the ischium and did not, apparently, observe that it 
was only present in the one sex. Kinghorn (1923 : 134), alone, seems to have 
observed that in Aprasia the spurs are present in males only. The present writer 
has confirmed that they are, likewise, secondary sex characters of the male in the 
other four genera mentioned above. 

Post-cloacal bones occur in male geckoes (Smith, 1933 : 9), a fact taken into con- 
sideration, with others, by McDowell & Bogert in aligning the Pygopodinae and 
Lialinae with the Gekkota ; but it seems unlikely that development of these bones 
to form external hooks would have occurred twice in Australasia, and nowhere else, 
pari passu with reduction of limbs, the acquisition of a serpentiform habit and 
sundry other features. So the position of Aprasia in the Pygopodidae seems 
assured. Within this family it is unique in its reduced dentition, degenerate eye, 
absent tympanum, rudimentary middle ear (Shute & Bellairs, 1953), the lack of a 
splenial bone (McDowell & Bogert, fig. 21) and vestigial shoulder girdle ; the latter 
is much more complete in both Pygopus and Lialis (Fiirbringer, 1870 : pi. i, figs. 
5, 6 : Kinghorn, 1926). It differs from Lialis in its paired parietal bones and 
resembles Pygopus, Paradelma, Delma and Pletholax in this respect. If, therefore, 
subdivision of the Pygopodidae into subfamilies is desirable, there would be 
justification for placing Aprasia in a distinct subfamily by itself. This procedure 
would, however, tend to obscure the fact that the genera Pygopus, Paradelma, 
Delma and Aprasia form an almost continuous linear series showing progressive 
limb-reduction and shortening and consolidation of the skull. Aprasia is the most 
completely " degenerate " of the series as it is also, probably, the most truly fossorial. 


As already mentioned, the material examined is divisible into five morphological 
groups which form a rational geographical picture, briefly as follows : 
I. A postocular shield present. 

(a) Snout prominent (fig. 3A), 1-5 to 2-1 times as long as the eye. 1 A suture normally 
runs from the nostril to the second labial, the nasal and second labial shields being 
in contact. 

(1) Presacral vertebrae 95-1 10 (mean 101-4). Mid -body scale rows 12 or 14. 
Victoria, South Australia, Nurina (W.A.). 

(1) Aprasia stviolata striolata. 

(2) Presacral vertebrae 88-94 (mean 91-8). Mid-body scale rows 12. Albany, 
Plantagent and Tambellup (W.A.). 

(2) Aprasia striolata glauerti. 
1 The method of measurement is important and is discussed below (p. 375). 



(b) Snout less prominent (Text-figs. 3B and 6), i-i to i-8 times as long as the eye. 
No suture running from the nostril. Presacral vertebrae 98-110 (mean 98-5) in 
males and 102-105 (mean 103-5) in females. Mid-body scale rows 14. Albany to 
Gerald ton (W.A.) : ? Northern Territory. 

(3) Aprasia pulchella. 

II. No postocular shield. Snout very prominent, 1-4 to 2-5 times as long as the eye (Text- 
figs. 3c and 7). A suture from the nostril reaches the prefrontal so that the nasal and 
second labial shields are not in contact or meet at a point only. 

(a) Presacral vertebrae 96-112 (mean 102-8) in males and 98-116 (mean 108) in 
females. Mid-body scale rows 12 (rarely 14). Albany to Shark Bay (W.A.). 

(4) Aprasia repens repens. 

(b) Presacral vertebrae 125, male and 137, female. Mid-body scale rows 14. Hermite 
Island (W.A.). 

(5) Aprasia repens rostrata. 

Fig. 3. Three male specimens of Aprasia from Albany, to show lateral head scales 
and proportions, a. Aprasia striolata glauerti subsp. nov. Holotype. b. Aprasia 
pulchella Gray. Perth Mus. 10955. c - Aprasia repens repens (Fry). Perth Mus. 

In this synopsis the assemblages have been given specific or subspecific status 
and justification of these treatments is called for. It will be observed that many 
of the characters used in the key " break down " if they are considered without 
reference to geographical distribution. For instance, mid-bod}/ scale rows and 


numbers of presacral vertebrae show a total range of variation which permits of no 
absolute differentiation between what are labelled as " species ", thus : 

A . striolata. 

A . pulchella. 

A . repen. 

Scale rows 

12- 14 


12- 14 





If, however, these characters are considered only in the geographical region where 
the three are sympatric, i.e. the Plantagenet Division of Western Australia, they 
provide in combination a clear cut differentiation, thus : 

A . striolata. . A . pulchella. A . repens. 

Scale rows 12 . 14 . 12 

Vertebrae . . 88-94 • 98-103 . 96-106 

A. pulchella is, additionally, completely differentiated from both of the others by 
its much less prominent snout with its less extensive rostral (less than one-third 
the diameter of the eye), the complete fusion of the nostril and first upper labial, 
and a distinctive colour pattern. Additional differences between A. striolata 
glauerti and A. repens repens exist in the same three features, viz. : Eye /rostral 
ratio 1-5-1-9 in glauerti vice 1-2-1-7 in repens, the suture behind the nostril running 
to the second labial in glauerti but to the prefrontal in repens and in colour. They 
are also completely differentiated by the post-ocular condition. Thus, the Aprasias 
of the Plantagenet Division can be subdivided into three different and distinct 
morphological groups which are not associated with age, sex, season, etc. It is 
reasonable to suppose that this fact signifies that there are three non-interbreeding 
assemblages and the only logical taxonomic treatment is to regard them as three 
distinct species. The most distinct morphologically is A. pulchella and this is 
also the species with the smallest geographical range (from Albany to Geraldton) 
and with the least variability. A . repens and A . striolata resemble each other more 
than either does A. pulchella and both show geographical variation of the same 

A. repens ranges northwards from Albany to the Monte Bello Islands and its 
distribution along this range may prove to be continuous, though the samples 
examined are not. The list of specimens examined, on pp. 382-383, is arranged in 
approximately south-north order, and it will be observed that, when due allowance 
is made for sexual and individual variation, there is a progressive increase in the 
number of presacral vertebrae from south to north ; there is also an increase in 
the number of mid-body scale rows from 12 to 14. It is, however, clear that the 
variation is not exactly correlated with latitude. For instance, in a series of sixteen 
males from Perth and its environs the vertebral range is from 95 to 108, whereas in a 
series from around Northam in almost the same latitude the range is noticeably higher, 
from 108 to 112. This and similar discrepancies may be due to chance and the 
short series available, but it seems at least equally possible that some environmental, 
possibly climatic, factor is involved. Although there is an overall increase in 
temperature from south to north, the isotherms do not follow an east-west course ; 



Northam has a more " continental " climate than Perth with higher temperatures 
in summer, when the isotherms are locally almost parallel with the coast (c.f. Year 
Book of Australia, 40, 1954, p. 33). Rainfall, too, though generally decreasing with 
decreasing latitude, also diminishes with distance from the coast. Unfortunately 
climatological information is not available for many of the localities from which 
specimens have been examined, and in any event the series are often very short. 
But if localities are grouped into regions around climatological stations it is possible 
to test, roughly, for correlation. The following table shows such groupings in 
relation to the six stations named. No climatic data being available for the Monte 
Bello Islands, the temperature and rainfall figures of Onslow have been used. 






.._,.„. ., A 




■ '** 






(° F.). 


96-0 (2) 





Katanning . 

97'3 (3) 

104-4 (8) 




Collie . 


103-0 (2) 




Perth . 

102 -6 (20) 

108-6 (25) 




Geraldton . 

107-0 (2) 

II2-0 (i) 

12/14 . 



York . 

110-7 (3) 

iii-o (2) 




Hermite Is. 

125-0 (1) 

I37-0 (I) 









here is no obvio 

us correlation between num 

bers of veri 

:ebrae and 

rainfall, either 

annual or monthly. But Text-fig. 4, in which vertebrae are plotted against the 
mean normal temperature for December, shows a correlation very clearly, though 
what its significance may be remains to be discovered. It is, however, difficult 
not to suspect that the correlation may be due to direct cause and effect, since it 
has been shown experimentally that the numbers of meristic structures in poikilo- 
thermic animals may be changed by varying the temperature during development. 
Wade Fox (1948), for example, has shown that in a Garter Snake, the numbers of mid- 
body scale rows, and ventral and subcaudal scutes are higher in groups of individuals 
whose mothers were kept at higher temperatures during pregnancy. It has also 
been established that in fishes (e.g. Taning, 1952, Lindsey, 1954) an increased 
temperature during the appropriate embryonic period will, in some species, result in 
an increased number of vertebrae, fin rays, etc. (and vice versa) within certain 
limits ; a decrease in temperature below a certain point may also, sometimes, 
result in an increase of the meristic characters. Detailed information regarding the 
breeding season of Aprasia in all areas, together with temperature records of the egg 
sites will be necessary to establish that we are, in fact, dealing here with a thermo- 
plastic effect. In the meantime all that can be said is that there is evidence of a 
cline correlated with the temperature in summer, a time of year when breeding 
may be expected to occur. There is a discontinuity in the series and it is, therefore, 
legitimate to subdivide it at the discontinuity, treating the two parts as subspecies. 
A. striolata, as already mentioned, also shows geographical variation in vertebral 
and mid-body scale numbers. It ranges from the south of Western Australia, 




§ * 
















o •» 




C 4> 
rt — 
rt O 





1, L_. 




- o 





CO ^ 

1 1 

Jh CD 

^ ft 

to S 

oJ fi 



CD 03 


ft -a 

>H *^ 


b0 co 

cd .y 



j) d.injEj^tlujsj 



where it is sympatric with A. repens in Albany, Plantagenet and Tambellup, across 
the south of the continent to western Victoria. In the west its vertebrae range 
from 88 to 94, there being no significant secondary sex difference, and the mid-body 
scales are constantly 12. In South Australia and Victoria the vertebrae range 
from 95 to no and 25% of the specimens seen have 14 scale rows. Again, as in 
A. repens, there appears to be some correlation with temperature, but not with 
rainfall, thus : 











(° F.). 

Albany . 

92-0 (11) 





9i-3 (3) 




Eyre (Nurina) 





Wudinna 1 






101 -4 (10) 

12 or 14 

21 -o 


1 Climatic data estimated as intermediate between Port Lincoln and Port Augusta. 

There is, however, inadequate information to establish the existence of tempera- 
ture plasticity and, moreover, a marked discontinuity becomes obvious when the 
series is plotted (Text-fig. 5). So in the present state of knowledge the recognition 
of two subspecies is indicated. 




• Eyres Penins. 
" Adelaide 

° O 

- Katanning 


- Eyre (Nurina) 


- Albany 


Subsp. glauertl 

1 1 1 

Subsp. itrlolata 
I > 1 1 









Fig. 5. Aprasia striolata. Average numbers of presacral vertebrae of local populations 
plotted against the mean December temperatures of the areas, 


Only sixteen specimens of A . pulchella with definite localities have been examined, 
all from points between Albany and Geraldton. Within this restricted area there 
appears to be little variation ; all the specimens have 14 scale-rows and the presacral 
vertebrae range between 96 and 98 in males (8) and from 102 to 105 in females (8). 
The type, a female without precise locality, has 104. The species may have a greater 
geographical range and there may be some geographical or temperature variation ; 
one specimen seen (Sydney, No. 12529) is an immature male with a much higher 
number of vertebrae, no. This specimen has no locality recorded against it in 
the registers of the Australian Museum, but formed part of a mixed collection 
mainly from the Northern Territory (Yirrikala Mission) and the northern parts of 
Western Australia. 

The number of scales longitudinally is associated with the number of vertebrae 
and consequently shows the same geographical variation within each species. There 
are, however, differences between the species in regard to the number of scales in 
relation to the number of vertebrae, i.e. relative size of the scales, and the differences 
appear to be significant. The ratio of the number of ventral scales to the number 
of vertebrae varies as follows : 

A. pulchella (18) 1*44 to 1-59. Mean 1-52 00-049. 
A.striolata (28) 1 -31 to 1 -50. ,, 1-4300-048. 
A.repens (48)1-23101-52. ,, 1-3500-06. 

Comparing the three on the criterion of the quotient of the difference of the means 
divided by the standard error of the difference of the means, the differences are all 
significant, thus : 

(a) A. pulchella j A. striolata d = 6-4. 


(b) A. striolata j A.repens ,, = 6-5. 

(c) A. pulchella I A.repens ,, = 11 «6. 

A. striolata is almost exactly intermediate between the other two, A. pulchella 
having the smallest scales. Increase in scale-size is often accompanied by complete 
or partial elimination of some scales. In Aprasia this effect is manifest in the head 
region. In A. repens, with the largest scales, the post-ocular has been eliminated 
and the pre-ocular is much reduced in size compared with the other two species 
(c.f. Text-figs. 3, 6 and 7). No variation in this character has been observed. 

The character of length of snout relative to size of eye can be appreciated by 
inspection, but is difficult to measure accurately. The method adopted was to 
measure the distance between the tip of the rostral and the anterior corner of the 
frontal as seen in plan projection on a plane parallel with that passing through 
the tip of the rostral and the centres of the eyes, using a camera lucida for the 
purpose. The eye diameter is that of the brill measured horizontally, i.e. excluding 
the ring of small scales that borders the " window ". The method cannot be claimed 
to give a high degree of accuracy, but it does provide directly comparable figures 


although the range of variation will almost certainly be exaggerated. Comparison 
between the subspecies shows no significant differences, thus : 

(a) A. striolata striolata (n). Mean i -67 o 0-14^! d = 1 -62. 
A. striolata glauerti (14). ,, 1-78 oo-i6j ^ 

(b) A. repens repens (65). ,, 1-96 o o-2o^\d = < 1, 

A . repens rostrata (2) . „ 2-1 ? J ad 

Between species, however, the differences are marked and significant, thus : 

A.pulchella (17). Mean 1-45 00-15. 
A. striolata (25). ,, 1-74,, 0-16. 

A. repens (67). ,, 1-96, ,0-20. 

(a) A. pulchellajA. striolata d = 6-o. 


(b) A. striolata I A. repens ,, = 5-5. 

(c) A. pulchella/A. repens ,, = 11-9. 

The character of the length of the rostral shield, as might be expected, is correlated 
with the length of the snout ; it shows no infraspecific variation that is certainly 
significant. As between species, however, the ratio of brill diameter to length of 
rostral* is : 

A. pulchella (17). Mean 3-90 o 0-53. 
A. striolata (25). „ i- 80 00-14. 

A. repens (65). ,, 1-5200-18. 

Comparing the differences of the means for significance : 

(a) A . pulchella I A . striolata d =16-2. 


(b) A. striolata J A. repens ,, = 8-8. 

(c) A. pulchella /A. repens ,, = 19-0. 

Thus, it seems to be established that infra-specific variation occurs mainly, if 
not exclusively, in meristic characters — the numbers of vertebrae and scales. Non- 
meristic characters such as the length of the snout relative to the eye (and the asso- 
ciated character of the relative proportions of the rostral shield), the presence or 
absence of a post-ocular shield, and relative scale size are significantly different 
between the samples that are believed to represent bio-species, but show no signi- 
ficant geographical variation within these groups. 


Aprasia striolata striolata Liitken 

Aprasia pulchella var. striolata Liitken, 1863, Vidensk. Medd. Kbh. 1862 : 300, pis. 1-2, figs. 3-3C 

Aprasia pulchella var. lineolata Liitken, 1863, loc. cit. 

Aprasia octolineata Peters, 1864, Monatsber. Ak. Berlin, 1863 : 233. 

Aprasia pulchella, Giinther, 1873 Ann. Mag. nut. Hist. (4) 12 : 145 (part). Giinther, 1875, Zool. 

* (Measured in plan projection.) 



Erebus and Terror, 2 : 10 (part). Boulenger, 1885, Cat. Lizards Brit. Mus. 1 : 246 (part). 

McCoy, 1888, Prodrom. Zool. Victoria, 17 : 233, pi. 161, fig. 1. Lucas & Frost, 1894, Proc. roy. 

Soc. Vict., n.s., 5 : 340. Lucas & le Souef, 1909, The Animals of Australia (Melbourne) : 219 

(part). Werner, 1912, Das Tierreich, 33 : 25 (part), fig. 5. Zietz, 1920, Rec. S. Aust. Mus. 

1 : 193 (part). Kinghorn, 1923, Rec. Aust. Mus. 14, 2 : 130 (part), figs. 7-9. Kinghorn, 

1926, Rec. Aust. Mus. 15 : 63, fig. 18. Kershaw, 1927, Vict. Nat. 43 : 341. Waite, 1929, 

Rept. and Amph. of South Australia (Adelaide) : 95, figs. 69, 70 
Ophiopsiseps nasutus (non Bocage), Jensen, 1901, Vidensk. Medd. Kbh. 1900 : 317, pi. 3. 
Ophioseps nasutus, {non Bocage), Fry, 1914, Rec. W . Aust. Mus. 1 : 181 (part). Kinghorn, 1923, 

Rec. Aust. Mus. 14, 2 : 128 (part), figs. 4-6. Kinghorn, 1926, op. cit., 15 : 62, fig. 17. 

Snout projecting and rounded, its length 1-5 to 2-0 times as long as the horizontal 
diameter of the brill (mean 1-67, cr 0-14). Rostral well visible from above, its 
length contained i-8 to 2-0 times in the length of the brill (mean 1*90, cr 0-07). A 
suture normally connects the nostril with the suture between the first and second 
labials so that the nasal and the second labial are in contact ; in one specimen 
only of the fifteen examined are these two shields separated. A small postocular 
separates the fourth labial from the supraocular. Premaxillary teeth may be 
present, though small, in females ; they are constantly present, and much larger, 
in males. Scales in 12 (80%) or 14 (20%) longitudinal rows at mid-body ; from 
140 to 155 scales, approximately, in a straight line from the mental shield to the 
vent in the proportion of 1*38 to 1-50 (mean 1*43, cr — 0-034, n. 13) per vertebra. 
Presacral vertebrae number 98 to 104 in males (5 counted) and 95 to no in the nine 
females. A well defined colour pattern of dark lines or rows of dots is usually 
present on the back. The largest female examined measures 133 mm. from snout 
to vent and the largest male 107 mm. 

This subspecies appears to range from western Victoria westwards to the Nurina 
Division of Western Australia. The reasons for its recognition have been discussed 
above. The material examined includes the immature specimens which Jensen 
misidentified as Ophioseps nasutus Bocage. The variation in the more important 
taxonomic characters is as follows : 

Verte- Snout. Eye 

Museum No. 


Cop. R.50.2 

Hochkirk, Vict. 

Sydney 11 759 • 

Jeparit, Vict. 

11760 . 

,, ,, 

B.M. 1902.7.30.8 

Renmark, S.A. 

,, 1902.5.30. 10 

>> >> 

,, 1902.5.30. 11 

y, >, 

,, 1902.5.30. 12 

,, ,, 

„ 1902.5.30. 13 

,, >, 

,, 1862.7. 19.4 

S. Australia 

,, 1862.7. 19.5 

,, ,, 

,, 1862.7. 19.6 

,, ,, 

Sydney 14382 . 

Wudinna, S.A. 

3466 . 




Perth R.5280 . 

Eyre, Nurina, W.A 









• ? 




• $ 



95 • 

• ? 




■ $ 




• ? 




• 6* 



98 . 

■ 6* 




■ 0* 




• ? 




• 6* 




■ <J 



99 • 

• ? 




• $ 




• ? 















Aprasia striolata glauerti subsp. nov. 
(Text-fig. 3A) 

Snout projecting and rounded (Text-fig. 3A), its length i-6 to 2-1 times as long 
as the horizontal diameter of the brill (mean 178, or = 0-16). The rostral is well 
visible from above, its length being contained 1-5 to 1-9 times in the length of the 
brill, the mean being 1-70 and <r 0-14. The condition of the nasal suture is similar 
to that of the eastern race, but two specimens of the fifteen are asymmetrical and 
have the nasal separated from the second labial on one side ; in one specimen the 
suture is incomplete and fails to reach either the interlabial or the naso-prefrontal 
suture. Mid-body scale rows are constantly 12 and there are from about 120 to 140 
from the mental shield to the vent in the proportion of 1-31 to 1-49 (mean 1-43, a* 0-058, 
n. 15) per vertebra. Presacral vertebrae number 88 to 93 in males (7) and 91 to 
94 in females (8). Premaxillary teeth present in both sexes, but smaller and some- 
times absent in females. Straw coloured above and beneath, sometimes with 
obscure darker markings suggesting dusky lines on the back. The largest male 
measures 109 mm. from snout to vent and the largest female 115 mm. 

The type series is from the " road districts " of Albany, Plantagenet and Tam- 
bellup, as listed below, the first mentioned being the holotype. 

Museum No. Locality. 




Perth 10949 





„ 6782 





„ 6782 





„ 6782 





„ 6782 





,, 6782 

' »» 




„ 6782 





„ 10950 





„ 1 095 1 





B.M. 1911.12. 





Perth 7214 





Sydney 12305 








> 88 







n 3468 

. [" Australia "] 






sia pw 


Ichella Gra 

3B and 6) 






1 • 



1 • 



1 • 



T • 



I • 



I • 



I * 



I • 



I • 



I • 


I • 



I ' 



I • 



I • 



Aprasia pulchella Gray, 1839, Ann. Nat. Hist. 2 ; 332. Gray, 1841, in Grey, Travels in Australia, 
2 : 428, 438, pi. 4, fig. 2. Gray, 1845, Cat. Lizards Brit. Mus. : 68 (part). Giinther, 1873, 
Ann. Mag. nat. Hist. (4) 12 : 145 (part). Giinther, 1875, Zool. Erebus & Terror, 2 : 10 (part). 
Boulenger, 1885, Cat. Lizards Brit. Mus. 1 : 246 (part). Lucas & le Souef, 1909, The Animals 
of Australia, (Melbourne) : 219 (part). Zietz, 1920, Rec. S. Aust. Mus. 1 : 193 (part). 
Kinghorn, 1923, Rec. Aust. Mus. 14, 2 : 130 (part). Kinghorn, 1926, Rec. Aust. Mus. 15 : 63 
(part). Loveridge, 1934, Bull. Mus. comp. Zool. Harv. 77 : 316. 



Ophioseps nasutus Bocage, 1873, /. Set. math. phys. nat. Lisboa, 4 : 232. Werner, 191 2, Das 
Tierreich : 33: 25. Fry, 1914, Rec. W. Aust. Mus. 1 : 181 (part). Kinghorn, 1923, Rec. Aust. 
Mus. 14, 2, 128 (part). 

Ophiopsiseps nasutus, Boulenger, 1887, Cat. Lizards Brit. Mus. 3 : 436. 

Aprasia brevirostris Werner, 1909, in Michaelsen & Hartmeyer, Die Fauna sudwest- Austral. 
2, 16 : 266, fig. 2. 

Snout slightly prominent and bluntly rounded (Text-figs. 3B and 6), its length 
i-i to i-8 times the length of the brill (mean 1-45, a 0-15). The portion of the 
rostral visible from above is less extensive than in any other species, its length 
being contained 3-1 to 5-0 times in the brillar length (mean 3-90, cr 0-53). No 
suture between the nasal and first labial shield, though in the holotype there is a 
vestige on the nasoprefrontal suture (Text-fig. 6). A postocular separates the 
fourth upper labial from the supraocular. Mid-bod}/ scale rows 14 ; from 145 
to 170 scales, approximately, on the mid-line between the mental shield and the 
vent in the proportion of 1-44 to 1-59 (mean 1-52, cr 0-049 > n - z &) P er vertebra. 
Presacral vertebrae 96 to no in males (9) and 102 to 105 in females (9) ; the possi- 
bility of geographical variation in this character has already been discussed. Pre- 
maxillary teeth absent in females. Pale brown above with indefinite dusky linear 
markings on the dorsal scales which can produce a lineolate appearance ; on the 
head the dark markings are more irregular and produce a vermiculate appearance. 
The largest male examined measures 108 mm. from snout to vent and the largest 
female 124 mm. 

This species is known with certainty to occur from Albany to Geraldton. 

Although Boulenger (1885) lists two specimens as " types " of A. pulchella, there 
is nothing in Gray's original description to indicate that he had more than one 
specimen whose dimensions are given as 2f in. from snout to vent with a tail of 
if in. Only one of Boulenger's specimens, the one listed below as B.M. 1946 . 8 . 30 . 93, 

Fig. 6. Aprasia pulchella Gray. Holotype female. 

3 8o 


approximates to these dimensions (snout to vent 2-8 in., tail 175 in.) and has ver- 
miculate head-markings (c.f. Text-fig. 6) as figured by Gray in 1841. The only 
doubts on the matter arise from the facts that (1) the original description states that 
the nostril lies " in the suture between the top of the first labial and anterior frontal 
plate " and (2) Gunther in 1873 stated that all the specimens then in the British 
Museum had only 12 mid-body scales. In the specimen now regarded as the type 
the first labial is fused with the shield above it and there are 14 scale rows. Gunther 
was patently in error, as was Boulenger (1885), also, when he failed to detect more 
than 12 scale rows and it seems likely that Gray, using only a hand lens, was misled 
by the vestige of a suture. 

Thanks to the courtesy of Professor Jorge the holotype of Ophioseps nasutus 
Bocage has been examined and compared with the other specimens listed. Unfor- 
tunately the specimen is in very poor condition ; it is in four fragments, breaks 
occurring after the ninth vertebra, in front of the nineteenth presacral, and imme- 
diately behind the pelvic girdle ; the tail tip is also wanting and the scales of the 
upper surface of the head are very badly abraded, so that their limits can only be 
determined with difficulty and a little uncertainty. It is also likely that the specimen 
has been partially desiccated at some time, since the cranium was full of air and 
the whole specimen measures only 73% of its pristine length (head and body frag- 
ments 95 vice 133 mm. and tail 38 vice 52 mm.). 1 Nevertheless, despite these 
defects, a sufficient number of characters can be deciphered or measured to make it 
virtually certain that the species involved is the same one which Gray had previously 
named Aprasia pulchella. Thus, for comparison with the characters of the species : 

0. nasutus 


Suture from nostril . 



Snout length /brill diam. 
Brill diam. /rostral length 



Mean 1-45 

„ 3'9 

Mid-body scales 


Mean 1 -52 

As noted above, the skull is visible dorsally and the ratio of frontal to nasal length 
(1*56) is similar to that in an example of A. pulchella (1*50). The presacral vertebrae 
number 89, but it is certain that a number have been lost from the ends of most 
of the fragments (one was found loose) and no significance can be attached to the 
number. The sex cannot be determined since, although no post-cloacal bones are 
present, there is a clear indication of the loss of at least 4 anterior caudal vertebrae. 
The first of the tail vertebrae has a well-defined fracture-plane through its centrum 
and so cannot be one of the most anterior caudal vertebrae which possess $0 
such split (Pratt : 184) ; in the numerous X-ray photographs of Aprasia that 
have been examined the fracture-plane is not evident anterior to the fifth to 
seventh caudal. 

1 The reduction cannot be due to the loss of a single large fragment of 52 mm. because when the tail 
and the head-plus-body are considered separately each is shorter than originally in the same proportion. 



The specimens examined and their characters are : 

Verte- Snout. 


Museum No. 


Perth 10954 • • • Albany 



» 2371 

Mt. Baker 







„ 3873 



Mt. Helena 





» 3342 


4629 A 


M.C.Z. 24460 

Mundaring Weir 

Perth 3409 . 

Upper Swan River 

B.M. — . 

Swan River 

M.C.Z. 24467 




B.M. 1946.8.30.9 


W. Australia 

Sydney 6553 




Mus. Bocage 

" Australia " 








1-6 . 

• 0" 



1-6 . 

• $ 



i-4 . 

• S 



i'5 • 

• o* 



1-6 . 

• 9 



1-4 . 

. 6* 



i'5 • 

• $ 



i-8 . 

• 0* 



i-5 • 

• 0* 



i-4 . 

• $ 



i-4 . 

• $ 



i-4 . 

• 6* 



i'3 • 

• ? 



i'5 • 

• $ 



1 -i 

• $ 



1-6 . 

• o*- 



i-5 • 

• ? 



I -2 




. i- 4 . 

• S 



1-4 . 


95 + 

89 + 

1-42 . 








Aprasia repens repens (Fry) 

(Text-fig. 3c) 

Aprasia pulchella (part), Gray, 1845, Cat. Lizards Brit. Mus. : 68. Giinther, 1875, Ann. Mag. 

nat. Hist. (4) 12 : 145. Giinther, 1875, Zool. Erebus & Terror, 2 : 10. Boulenger, 1885, Cat. 

Lizards Brit. Mus. 1 : 246. Werner, 1909, in Michaelsen & Hartmeyer; Die Fauna siidwest 

Austral. 2, 16 : 266. Lucas & le Souef, 1909, The Animals of Australia (Melbourne) : 219. 

Werner, 1912, Das Tierreich, 33 : 25. Zietz, 1920, Rec. S. Aust. Mus. 1 : 193. 
Ophioseps repens Fry, 191 2, Rec. W. Aust. Mus. 1 : 178, fig. 
Aprasia repens, Kinghorn, 1923, Rec. Aust. Mus. 14, 2 : 132, figs. 10-12. Kinghorn, 1926, 

Rec. Aust. Mus. 15 : 64. Loveridge, 1934, Bull. Mus. comp. Zool. Harv. 77 : 317. 
Lialis repens, Kinghorn, 1926. torn. cit. fig. 19. 

Snout strongly projecting (Text-fig. 3c), its length 1-4 to 2-5 times as long as the 
brill (mean 1-96, cr 0-20). Rostral well visible from above, its length contained 
i-o to 2-4 times in the brillar diameter (mean 1-52, a 0-18). A suture connects the 
nostril with the anterior border of the prefrontal so that the first labial and the 
prefrontal are in contact ; in two specimens the four shields (nasal, prefrontal, 
first labial, second labial) meet at a point and in another example the two sides arc 
not symmetrical the nasal being in contact with the second labial on the right 
side but separated from it on the left. There is no post ocular, the fourth labial 
reaching the supraocular. Premaxillary teeth absent in females. Scales in 12 
(75 specimens) or 14 (1 specimen) rows at mid-body and between 125 and 165, 
approximately in a median line from the mental shield to the vent in the proportion 
of 1-23 to 1-52 (mean 1-35 ; cr 0-06 ; n. 46) per vertebra. Presacral vertebrae 



96-112 in males and 98-116 in females, there being geographical, possibly temperature 
correlated, variation as discussed above. The ground colour is pale straw, above 
and below, often with ill-defined dark longitudinal lines or rows of dots on the back 
and especially on the flanks ; head sometimes with dark markings ; upper lip and 
throat chrome yellow ; tail red or yellow. The largest male examined is 150 mm. 
from snout to vent and the largest female 165 mm. 

The subspecies ranges from Albany to Victoria (W.A.) and possibly further 
north. Werner (1909) records a specimen in the Perth Museum as from Shark Bay, 
but there is no present record of such a specimen. The only possibility is specimen 
R.i 1360, a male which bears a label " M & H " (= Michaelsen and Hartmeyer) ; 
but Werner's record is not listed as having been obtained at any of the stations 
where Michaelsen and Hartmeyer worked and which are all numbered. In addition 
its vertebral count (104) is much more compatible with an origin near Perth. 

The material examined contained these specimens : 



Museum No. 







Perth 10952 . . . Albany 






„ io953 







B.M. 1931.7.1.139 

Mt. Toolbrunup 






Perth 2868 







„ 2869 






„ 2870 







„ 2871 





















.. 433^ 







.. 433B 







» 373 • 







Sydney 8015 

y » 






Perth 10246 . . . Williams 






10322 . . . Bunbury 






10895 . . West Midland 







,, ,y 






„ 10893 

y> »> 






» 4173 







„ 766 . 

Midland Junction 






„ 3734 

Rottnest Isld. 






„ 3759 

>, 11 






» 3765 

.» y, 






» 4582 

y, ,y 






M.C.Z. 33027 







33° 28 























? • 







Museum No. 
Perth 4373 
Perth 3347 

„ 2596 

.. 1742 

,, 6944 

„ 631 
Perth 9664 

„ 9665 

„ 2837 

.. 8745 

„ 8185 

„ 9093 

„ 9094 

>> 5344 

M 123 • 
» 7405 
„ 74O6 
M 7390 
„ 53IO 

„ 8721 

» 1374 

„ 5351 

» 2397 

„ 4186 

» 707 • 

» 4383 

M.C.Z. 24427 
Perth 4356 

„ 4357 
Sydney 9138 

Perth 5492 
„ 8891 

» 4913 

„ 6927 


„ 1730 
„ 5064 
M.C.Z. 24458 

Perth 1 1360 


East Perth 



Gooseberry Hill 







Mt. Lawley 

E. Freemantle 

Balcatta Beach 


Bakers Hill 

Newmarracarra (Vict. 

Eradu (Vict.) 


W. Australia 





























































— • 105 



















2 -I 
2 -O 







2 -o 




* This specimen is ovigerous. 



Aprasia repens rostrata subsp. nov. 

(Text-fig. 7) 
Aprasia sp. Hill, 1955, Proc. Linn., Lond. 165, 2 : 115. 

Holotype a male, collected 17.viii.52 on Hermite Island and allotype a female 
collected on the same island 22. v. 52. 

Snout very strongly projecting (Text-fig. 7) its length 2-0 times and 2-2 times as 
long as the diameter of the brill in male and female respectively. Rostral shield 
drawn out posteriorly so that the length of the portion visible from above is contained 
0-9 (<?) and o-8 (?) times in the brillar length. The suture running backwards from 
the nostril meets the suture between the first and second labials in such a position 

Fig. 7. Aprasia repens rostrata subsp. nov. Allotype female. 

that these two labials, the nasal and the prefrontal meet at a point. No post- 
ocular shield, the supraocular produced ventrally to make contact with the fourth 
labial. Premaxilliary teeth present in the male but absent in the female. Scales 
in 14 longitudinal rows at mid-body. Scales on the mid-line between the mental 
shield and the vent are about 175 in the male and 190 in the female, counted to the 
nearest five. Presacral vertebrae 125 in the male and 137 in the female. 

Pale brown above and white beneath. A broad darker lateral band on the sides 
of the neck which breaks up posteriorly into three fine brown lines. Three additional 
fine brown lines on the nape, but the mid-dorsal region of the body not lined ; a 
single mid-dorsal dark line along the tail. Another brown line on each side of the 
belly. Head vermiculate. 

Dimensions : 

cJ. Snout to vent 99 mm. ; tail 65 mm. (incomplete). 
?. „ >, » 109 mm. ; „ 13 mm. ( „ ). 



Thanks are due to Sir William Penney who, despite his many preoccupations 
with the major objects of the expedition to the Monte Bello Islands, arranged for 
collections of the fauna and flora of the islands to be made, and to Surgeon 
Commander G. Wedd, R.N., who organized and supervised these activities. The 
resulting material touched off the present investigation, but no progress could have 
been made without the assistance of Mr. L. Glauert, Mr. J. R. Kinghorn, Mr. F. W. 
Braestrup and Mr. A. Loveridge, who have been most generous in lending comparative 
material from their museums at Perth, Sydney, Copenhagen and Harvard. The 
author is also grateful to his colleagues Mr. N. B. Marshall, and Mr. D. W. 
Tucker, for help with ichthyological references, and to Mr. P. E. Purves for the 
large number of X-ray photographs that had to be taken. 

Boulenger, G. A. 1 885. Catalogue of Lizards in the British Museum {Natural History). 

Second edition. 
Fox, Wade. 1948. Effect of Temperature on Development of Scutellation in the Garter 

Snake, Thamnophis elegans atratus. Copeia, 1948 : 252-266. 
Furbringer, M. 1870. Die Knochen unci Muskeln der Extremitdten bei den Schlangendhnlichen 

Sauriern. Leipzig, 136 pp., 23 pis. 
Gray, J. E. 1841. A Catalogue of the Species of Reptiles and Amphibia, hitherto described 

as inhabiting Australia, with a description of some New Species from Western Australia 

and some remarks on their geographical distribution. Appendix E. to Grey, G., Journals 

of two Expeditions' of Discovery in North-West and Western Australia, 2 : 422-449. 
Gunther, A. 1873. Notes on, and descriptions of, some Lizards with Rudimentary Limbs 

in the British Museum. Ann. Mag. nat. Hist. (4), 12 : 145-148. 
Jensen, A. S. 1901. Om Ophiopsiseps nasutus du Bocage, etc. Vidensk. Medd. naturh. 

Foren. Kbh. 1900 : 317-328, pi. 3. 
Kinghorn, J. R. 1923. Studies in Australian Reptiles, No. 3 : On the genera and species 

of Ophioseps and Aprasia. Rec. Aust. Mus. 14, 2 : 126-134. 

1926. A brief review of the family Pygopodidae. Rec. Aust. Mus. 15 : 40-64. 

Lindsey, C. C. 1954. Temperature controlled meristic variation in the Paradise Fish 

Macropodus opercularis (L.). Canad. J. Zool. 32 : 87-98. 
Loveridge, A. 1934. Australian Reptiles in the Museum of Comparative Zoology, Cam- 
bridge, Mass. Bull. Mus. comp. Zool. Harv. 77 : 316-317. 
Lutken, C. 1863. Nogle nye Krybdyr og Padder. Vidensk. Medd. naturh. Foren. Kbh. 

1862 : 292-311, pis. 1-2. 
McDowell, S. B. and Bogert, C. M. 1954. The Systematic Position of Lanthanotus and 

the Affinities of the Anguinomorphan Lizards. Bull. Amer. Mus. nat. Hist. 105 : 1. 
Pratt, C. W. M. 1946. The plane of fracture of the caudal vertebrae of certain Lacertilians. 

/. Anat., Fond. 30 : 184-188. 
Shute, C. C. D. and Bellairs, A. d'A. 1953. The cochlear apparatus of Geckonidae and 

Pygopodidae and its bearing on the affinities of these groups of lizards. Proc. Zool. Soc. 

Lond. 123 : 695-709, 2 pis. 
Smith, M. A. 1933. Remarks on some Old World Geckoes. Rec. Indian Mus. 35 : 9-19. 
Vedel Taning, A. 1952. Experimental study of meristic characters of fishes. Biol. Rev. 

27 : 169-193. 
Werner, F. 1909. Reptilia exkl. Geckonidae und Scincidae. Fauna Stidw est- Aust. 2, 16, 

251-278, pis. 14-15. 

191 2. Eublepharidae, Uroplatidae, Pygopodidae. Fierreich, 33 : ix + 33. 


S APR 1956 




R. w. SIMS 

ZOOLOGY Vol. 3 No. 10 

LONDON : 1956 





Pp. 387-438 ; Plates 13-14 ; 2 Text-figures 



ZOOLOGY Vol. 3 No. 10 

LONDON: 1956 

(NATURAL HISTORY), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical Series. 

Parts will appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

This paper is Vol. 3, No. 10 of the Zoological series. 


Issued March, 1956 Price Fifteen Shillings 




By R. W. SIMS 
(with Field Notes and Photographs by W. T. Loke) 

(Received 13th July, 1955) 



Introduction ........... 389 

Collecting Localities ......... 391 

Annotated List of Birds Collected in the Wahgi Region by Mr. F. 

Shaw-mayer .......... 393 

References ........... 437 


This is a report on the birds collected by Mr. F. Shaw-Mayer in the Central Highlands of 
New Guinea, mainly in 1950 and 1951. Two-hundred and seventy- two specimens were 
obtained in the Wahgi region, principally on Mount Giluwe. Eighty-eight species and races 
are represented of which one race was new to science. 


i. General 

Mr. F. Shaw-Mayer has lived and collected for many years in New Guinea and has 
travelled widely in the eastern half of the country. In recent years he has travelled 
mainly in the Central Highlands and a collection of birds from that area forms the 
subject of this report. 

The Central Highlands of New Guinea were almost inaccessible prior to 1945 and 
little was known of the region, its inhabitants or its fauna. In recent years, however, 
the establishment of an airfield at Nondugl has made it more accessible and advantage 
has been taken of this for making collections by others besides Mr. Shaw-Mayer. In 
1950, and again in 1952, Mr. E. T. Gilliard of the American Museum of Natural 
History made large collections of birds in the Wahgi region ; and during the inter- 
vening year, 195 1, Count Gyldenstolpe also visited the area. The reports on these 

ZOOL. 3, IO. 25 



expeditions were in the course of preparation when I started identifying and making 
notes on Mr. Shaw-Mayer's specimens. The appearance of the first (Mayr and Gilliard, 
1954) greatly facilitated the completion of these notes. The facts recorded below and 
the taxonomic conclusions arrived at differ for the most part only in points of detail, 
such as ranges of dimensions and various minor aspects of plumage colour, differences 
which one would expect to find in two sets of random samples from the same area. 
There are, however, a number of additional facts some of which suggest slight 
modification to Mayr and Gilliard's conclusions. These notes had been completed 
when I learnt that Count Gyldenstolpe's report was in the press. When it was 
published (Gyldenstolpe, 1955) I read it in conjunction with my own notes and 
referred to many specimens again. Generally there was a large measure of agreement 
but on one or two points our conclusions were not identical, mainly because of 
differences in age or sex of the specimens, or as a result of making comparisons with 
material from different areas. 

Mr. Shaw-Mayer gave brief field notes on some specimens and a number of these 
are quoted here. Additional important field notes were made by Mr. Loke Wan Tho of 
Singapore. Mr. Loke joined Mr. Shaw-Mayer during September and October, 1952, 
when they travelled together up the Wahgi valley from Nondugl to Tomba. There 
Mr. Loke compijed most of his field notes which he kindly sent to me ; and extracts 
from most of them are included here. Two of his photographs are also published with 
his kind permission. 

I wish to acknowledge the generous co-operation of the authorities in the museums 
at Berlin, Harvard, Leiden, New York and Pittsburgh. They have very kindly lent 
me specimens for examination or made critical notes on specimens sent to them for 
comparison. I would also like to acknowledge the help afforded to me by Mr. R. P. 
Derek Goodwin. He made a preliminary examination of the Shaw-Mayer Collection 
and I found his notes most useful. Finally I wish to thank Mr. J. D. Macdonald for 
reading the manuscript and for his helpful criticism. 

2. Methods 

The report of the American Museum of Natural History Expeditions (Mayr and 
Gilliard, 1954) contained descriptions of the topography and ecology of the area, and 
lists of the birds typical of the various habitats and their zoogeographical affinities. 
These subjects, therefore, receive no further treatment in this report except where 
new information is presented. 

The sequence of the families, genera and species and the nomenclature are that of 
Mayr (19416). In that work a synonymy of each name is given and consequently 
only the original reference to each name is normally given here. 

The specimens are listed under their British Museum (Natural History) register 
numbers together with information provided by the collector. The dimensions are 
given in millimetres ; the wings were flattened when they were measured so the 
measurement given is the arc ; the bill measurement is that of the exposed part of 
the culmen. 




3. Summary of New Information 
[a) Additions to the list of the birds of the Wahgi region (Gyldenstolpe, 1955). 

Casaurius bennetti shawmayeri. 
Dupetor flavicollis gouldi. 
Falco peregrinus ernsti. 
Porzana pusilla palustris. 

Alcyone azurea ochrogaster. 
Cisticola exilis diminuta. 
Petroica bivittata bivittata. 
Aplonis metallica metallica. 

Neositta papuensis alba. 

(b) Zoogeographical affinities of Wahgi region birds not listed by Mayr and Gilliard, 


Southern affinities (Australia) : South-eastern affinities : 

Porzana pusilla palustris Casuarius bennetti shawmayeri (Kratke 

Cisticola exilis diminuta Circus spilonotus spilothorax 

Petroica bivittata bivittata 

North-eastern affinities (Huon Peninsula) : 

Turnix maculosa giluwensis 

Northern affinities (Sepik Region) 
Alcyone azurea ochrogaster 
Astrapia stephaniae femina 

Western affinities (Mountains of Dutch New 
Guinea) : 
Neositta papuensis alba 

(c) Habitats of Wahgi region birds not listed by Mayr and Gilliard, 1954. 

Grasslands — 4,000 ft. to 7,500 ft. Mid-mountain forest — 7,500 ft. to 

9,000 ft. 

Dupetor flavicollis gouldi Casuarius bennetti shawmayeri 

Anas super ciliosa rogersi Scolopax saturata rosenbergii 

Circus spilonotus spilothorax Aplonis metallica metallica 

Turnix maculosa giluwensis Astrapia stephaniae femina 

A ley one azurea ochrogaster Neositta papuensis alba 
Cisticola exilis diminuta 

Timber-line, edge of the alpine grasslands — 11,000 ft. 

Petroica bivittata bivittata 


October, 1946 : Menebe, Mount Hagen, 5,000 ft. 

October, 1950 : Headwaters of the Minj River, Kubor Range, 8,000 to 9,000 ft. 

November, 1950 ; Minj, Wahgi River, 5000 ft. ; and the forested slopes of the 

Kubor Range. 
December, 1950 and January, 195 1 ; Forested ridges on the west slopes of the 

Hagen Range near Welya, 7,500 ft. to 10,000 ft. 

1 Racial identity problematical, Gyldenstolpe (1955). 

2 Possibly includes Neositta papuensis wahgiensis Gyldenstolpe, 1955. 




February and March, 1951 : South slopes of the Hagen Range near Tomba, 8,000 

to 10,000 ft. 
March to July, 1951 : Plateau grasslands north of Mount Giluwe, 7,300 ft. Lamende 
Range north-west of Mount Giluwe, 8,500 ft. (23rd March). Moss Forest, 
10,000 ft.; and the alpine grasslands, 11,000 ft.; Mount Giluwe (24~28th 

Moss Forest, 10,000 ft.; and the alpine grasslands, 11,000 feet, Lamende 
Range (nth to 14th May). 

Forested north slopes, 7,500 to 8,500 ft.; and the mid-mountain grasslands, 
9,000 ft., Mount Giluwe (20th to 30th May). 
Mount Giluwe rises to 13,414 ft. above sea-level from a grassland plateau of 7,300 ft.. 
The lower slopes of the mountain are forested up to about 9,000 ft. where the mid- 
mountain grasslands commence. These are separated from the alpine grasslands of 
the summit by the moss forest that extends approximately from 10,000 ft. to 11,000 
ft.. The distribution of the forests and grasslands appears to be similar on the Lamende 
Range and characteristic of the mountains of this area ; fuller details, particularly 
of the Bismarck, Hagen and Kubor Ranges, were given by Mayr and Gilliard (1954 : 


Casaurius bennetti shawmayeri Rothschild 

Casuarius papuanus shawmayeri Rothschild, 1937, Bull. Brit. Orn. Club, 57, p. 120. Arau 
district, Kratke Mountains, New Guinea. 

1954.17.266 ; i $ ; Menebe, north of Mount Hagen, 5,000 ft.; October, 1946. 

Measurements, i ? — height of back from ground, 860 ; total length, 1,170. 
Colours : bill and casque, shiny black ; legs, horn ; iris, brown. Stomach Contents : 
remains of hard fruits. 

The specimen and a drawing of the head and neck at death agree with the type. The 
record extends the known range of shawmayeri about 150 miles north-west from the 
Kratke Mountains to the Hagen Range. It is probable that this race is found through- 
out the Wahgi Valley area, but it remains to be seen how far it extends into the 
mountains on each side. Mr. Gilliard saw a cassowary on Mount O'-Mar in the Kubor 
range which he believed was this race (Mayr and Gilliard, 1954 : 331), and the present 
specimen taken near Mount Hagen supports his provisional identification. 

Dupetor flavicollis gouldi (Bonaparte) 
Ardetta gouldi Bonaparte, 1855, Consp. Gen. Av. 2, p. 132. New South Wales, Australia. 

1953. 17. i ; 1 (J ; Mount Giluwe, 7,300 ft.; June, 1951. 

Measurements : 1 <$ — wing, 209 ; tail, 76 ; bill, 69. Colours ; bill, upper 
mandible dark horn and lower mandible light horn ; legs, brown-grey ; iris, yellow. 
Stomach Contents : water insects, 


Anas superciliosa rogersi Mathews 

Anas superciliosa rogersi Mathews, 1912, Austr. Av. Rec. 1, p. 33. Augusta, south-western 

1953. 17.2-4 ; i (J, 2 juv. cJ ; Mount Giluwe, 7,300 ft.; June, July, 1951. 

Measurements : 2 juv. (J — wing, 229, 239 ; tail, 68, 71 ; bill 46, 48. 1 <J — wing 
250 ; tail 96 ; bill 47. Colours : bill, slate (ad. <J black at base of lower mandible) ; 
legs, pale brown flesh (ad. £ greenish pale brown, darker on webs of feet) ; iris, 
brown. Stomach Contents : insect larvae, grass seeds and river gravel. 

These birds were taken on the plateau north of Mount Giluwe where the species 
was plentiful. The June birds appear to be juveniles because the feathers of the 
undersurface have a dark brown streak unlike the adult, where they are dark brown 
and fringed with cinnamon. The primaries are fully grown and show no signs of wear, 
yet the wing is shorter than the adult's. In this respect the young birds approach the 
smaller size of the other New Guinea race, pelewensis, which differs from rogersi only 
in the shorter wing — -pelewensis, 230 ; rogersi ,250 — Amadon (1945 : 3) ; nevertheless, 
this pair appear to be young birds of the larger form, rogersi. I have not seen 
any material of comparable age and plumage, but it is likely that the primaries of 
the juvenile plumage which persist throughout the first adult plumage are shorter 
than the primaries of the full adult plumage. The young birds have started to moult 
and a few adult patterned feathers have come in on the undersurface ; the wing does 
not appear to be involved in the moult. The July bird, taken nearly six weeks later, 
is completing moult. The feathers of the undersurface of all three specimens are 
variably stained a rust colour, probably with iron oxide. 

Salvadorina waigiuensis Rothschild and Hartert 

Salvadorina waigiuensis Rothschild and Hartet, 1894, Novit. Zool. 1, p. 683. ? Waigeu, possibly 
Vogelkop, north western New Guinea. 

1953. 17.5-6 ; 2 ? ; Mount Giluwe, 7,300-7,200 ft.; April, July, 1951. 

Measurements : 2 °- — wing, 180-185 ; tail, 95 ; bill, 34-35. Colours : bill, dull 
yellow with black along the centre of the upper mandible ; legs, yellowish horn (webs 
of feet almost black) ; iris, dark brown. Stomach Contents : river gravel, water 
beetles and other small aquatic insects. 

Accipiter melanochlamys schistacinus (Rothschild and Hartert) 

Astur melanochlamys schistacinus Rothschild and Hartert, 191 3, Novit. Zool. 20, p. 482. Mt. 
Goliath, Oranje range, central New Guinea. 

1953. 17.7-8 ; i (J, i $ ; Mount Giluwe ; 8,000 ft.; May, July, 1951. 
Measurements : 1 <J — wing, 216 ; tail, 160 ; bill, 15-5. 1 $ — wing, 251 ; tail, 
184 ; bill, 18. Colours : bill, black (cere — $ yellow and $ orange-yellow) ; legs, 


J bright yellow and ? orange ; claws, black ; iris, <J bright yellow and $ bright orange. 
Stomach contents : £, remains of small bird ; ?, remains of frog. 

The female (July) is in moult and the new black feathers of the back have a 
greenish metallic lustre in contrast to the worn feathers which have faded to a rusty 
brown-black. The male (May) is a young bird in fresh immature plumage. The 
exposed parts of the feathers of the back are chestnut in colour edged with rufous brown, 
the unexposed parts being white. The feathers of the shoulders and most of the head 
are white tipped with black, while the crown is chestnut in colour. The primaries and 
the retrices are dark brown with black-brown barring above ; they are silvery below 
with irregular cinnamon coloured areas, particularly on the inner webs. The feathers 
of the under-surface are cream in colour with a median chestnut stripe while those of 
the legs are more buffy with a narrow rufous stripe. 

Circus spilonotus spilothorax Salvadori and D'Albertis 

Circus spilothorax Salvadori and D'Albertis, 1875, Ann. Mus. Civ. Genova, 7, p. 807. Yule 
Island, south-eastern New Guinea. 

I 953-i7-9 ; 1 c? ; Mount Giluwe, 7,300 ft.; July, 1954. 

Measurements : 1 $ — wing, 375 ; tail, 220 ; bill, 21-5. Colours : bill, black 
(cere, bluish horn ; dorsal surface, greenish yellow.) ; legs, dull yellow ; claws, 
black ; iris, bright yellow ; inside of mouth, blue-grey. 

" I often observed this fine bird soaring over the grasslands of the plateau, some- 
times high in the air, at other times a few feet above the grass. Several times it came 
to rest on the ground, but it was not observed to be eating anything. At the approach 
of the hunter it was off, sailing with speed in ever widening circles. Once noticed at 
dusk, but more often during the early morning. It appeared to be alone." (F. S.-M.). 

In comparison with other birds of this race this specimen is considerably darker in 
colour, both above and below. The upper-parts are a rusty black and lack the typical 
silvery white wash which is present only over the upperside of all except the inner 
and outer pairs of rectrices. The inner rectrices are almost black above and the outer 
a faded chestnut colour, none being spotted like those of other specimens. This bird 
exhibits a type of melanism not uncommon among birds of prey, but it appears to be 
darker than any reported by Rand (1941a : 1). 

Falco peregrinus ernesti Sharpe 
Falco ernesti Sharpe, 1894, Ibis, Ser. 6, 4 : 545. Mt. Dulit, Sarawak. 

^S-^-io-; icJ; Mount Giluwe, 7,3000 ft.; July, 195 1. 

Measurements : 1 <$ — wing, 292 ; tail, 144 ; bill, 19. Colours : bill, dark horn 
around cere and gape, base of mandible yellow ; legs, yellow (claws, black) ; iris, 
black (skin around eye, yellow). 



Aepypodius arfakianus (Salvadori) 

Talegallus arfakianus Salvadori, 1877, Ann. Mus. Civ. Genova, 9 : 333. Arfak Mountains, north- 
western New Guinea. 

1953.17.11-12 ; i (J, i ? ; Mount Giluwe, 9,000 ft.; April, July, 1951. 

Measurements : 1 <J — wing, 278 ; tail, 132 ; bill, 31. 1 ? — wing, 272 ; tail, 145 ; 
bill, 30. Colours : bill, yellowish-green-horn ; legs, greenish-yellow ($, frontal 
scutes, dark horn) ; iris, greenish yellow ; head, pale bluish white, reddish colouring 
along top of gape from back of beak to below eye (<?, purple comb and wattles, 
bluish area on crown to hinder end of the base of the comb ; $, two small pinkish 
wattles — no comb — bluish area above wattles). Stomach contents : pieces of hard 

The birds were taken in the forest on the north slopes of Mount Giluwe. An egg 
was found in the oviduct of the female (July). 

Synoicus ypsilophorus lamonti Mayr and Gilliard 

Synoicus ypsilophorus lamonti Mayr and Gilliard, 195 1, Amer. Mus. Novit. No. 1524, p. 1. 
Central Highlands, New Guinea. 


1953 . 17 . 13-24 ; 7 c?, 5 ? ; Mount Giluwe, 7,500-7,200 ft.; April, May, 1951. 

Measurements : 7 $ : wing, 90-100 ; bill, 12-13-5. 5 ? — wing, 93-98 ; bill, 12-13. 
Colours ; bill, black, bluish horn at base ; legs, greenish yellow ; iris, yellow-orange. 
Stomach contents : chiefly grass seeds, some insects and gravel. 

Most of the specimens of this fine series match closely 2 (J and 2 $ of the race 
lamonti taken by Mr. Gilliard at Tomba, which is only a few miles north of Mount 
Giluwe. The exception is the male bird (No. 1953. 17. 19) taken on 27th May. In 
general colour it is a much paler brown, more heavily barred above and below and the 
white shaft streaks of the feathers of the back are wider and more pronounced. It 
was sent to the American Museum of Natural History where, in Mr. Gilliard's absence, 
Dr. Amadon kindly compared it with other material of 5. ypsilophorus. He found 
that it was unlike any other bird of this species and that the differences between this 
specimen and other examples of lamonti are greater than those between lamonti and 
neighbouring races. As these races appear to be fairly stable in their characteristics 
we have concluded that this specimen is probably aberrant. 

" This quail was plentiful on the grasslands at 7,000 feet, and on the grasslands on 
the north slopes of Mount Giluwe up to 8,000 feet. Sometimes a single bird would flush, 
at other times three or more. Much of the country is peaty, almost swampy, and these 
conditions seem to suit it best " (F. S.-M.). 

Field Notes. Six of these quail were started up in pairs from the long grass near 
Nondugl. A roughly made nest of grass was found on the ground among the weeds 
near some land under cultivation on 15th September. The nest contained four 
buff-coloured eggs with thickly distributed minute brown spots. 


Excalfactoria chinensis subspecies ? 
Tetrao chinensis Linnaeus, 1766, Syst. Nat. 12th ed., 1, p. 277. China. 

1953.17.25-33 ; 3 cj, 6 ? ; Mount Giluwe, 7,500-7,200 ft.; April, May, 1951. 

Measurements : 3 <J — wing, 65-68 ; bill, 8-9. 6 $■ — wing, 65-70 ; bill, 9-10. 
Colours : bill, black ; legs, orange, orange-yellow (2 $ with greenish tinge) ; iris, 
3 c? — brownish-red ; 6 ? — dark brown. Stomach Contents : grass seeds and gravel. 

The series was collected on the grasslands north of Mount Giluwe. The birds are in 
fresh plumage with the exception of the female taken on 7th April which is in moult 
although it had an egg in the oviduct. Two races of E. chinensis have been described 
from New Guinea, papuensis Mayr and Rand from Mafulu in the south-east and 
novaeguineae Rand from the Balim River in the Snow Mountains. Mount Giluwe is 
approximately midway between the type localities of these races and, as might be 
expected, the birds from there exhibit characteristics of both. The series was com- 
pared with specimens of both races collected by Dr. Rand. The adult males differ 
but little from a male taken at Lake Daviumbo on the Fly River and compared with 
typical papuensis the bluish colour of the head is slightly less pronounced and the 
small feathers of the head have dark grey-brown tips ; in this respect they approach 
the characteristics of novaeguineae. The adult female from Mount Giluwe differ 
from papuensis only in the colour of the under-parts, where there are gradations from 
the richer colour of papuensis to the paler colour of novaeguineae. 

The series is almost intermediate between papuensis and novaeguineae. I am 
not, therefore, naming them racially. Furthermore, Dr. Rand's birds are in old 
plumage, whereas in most of these the plumage is fresh. 

Turnix maculosa gilutvensis Sims 

Turnix maculosa giluwensis Sims, 1954, Bull. Brit. Orn. Club, 74, p. 37. Mount Giluwe, Central 
Highlands, New Guinea (8,000 ft.). 

I 953-i7-34-4i 1 ; i(J,i juv. $; 3 <j>, 3 juv. $ ; Mount Giluwe, 8,000-7,200 ft.; 
May, June, 1951. 

Measurements: 2 $ — wing, 65; tarsus, 18. 5-19; bill, 10-n. 6 ? — wing, 
70-75 ; tarsus, 19-20 ; bill, 11-5-14. Colours : bill, yellowish, ridge and tip dark 
horn; feet, yellow with greenish tinge; iris, cream. Stomach contents: grass 
seeds and small insects. 

This fine series of Button-quail was taken on the open and lightly wooded grass- 
lands around Mount Giluwe. It appears that this is the first time that such a large 
series of T. maculosa has been taken in one locality in New Guinea ; the males, 
particularly, are rare to collections. On examination these birds were found to be 
distinct enough to recognize as representatives of a new race of T. maculosa in 

1 Type 1953.17.40 


comparison with the other two New Guinea races, furva Parkes from Finschhafen 
on the east coast of the Huon Peninsula and horsbrughi from Yule Island off the 
south-east coast. They differ from these races by the underparts being considerably 
paler, but resemble furva by being very dark above. In size giluwensis is closer to 
horsbrughi than to furva, which is slightly larger. The one June bird is in fresh 
plumage the others are involved in moult. 

Rallus pectoralis captus Mayr and Gilliard 

Rallus pectoralis captus Mayr and Gilliard, 1951, Amer. Mus. Novit. No. 1524, p. 2. Tomba, 
Central Highlands, New Guinea. 

1953.17.42 ; i(J; Hagen Range, 8,000 ft.; February, 1951. 

1953.17.43-45 ; 1 (J, I juv. (?, 1 $ ; Mount Giluwe, 7,500-7,300 ft.; April, June, 

Measurements : 3 <J — wing, 102-104 ; tail, 37-40 ; bill, 33. 1 $ — wing, 102 ; 
tail, 40 ; bill, 34. Colours : bill, dull red, dark horn near tip ; legs, brownish grey ; 
iris, yellow to brown (juv. <J, grey brown). Stomach contents : slugs and water 

The adult male and female taken in February and April respectively are in worn 
plumage ; the barring of the abdomen is faded and the flanks are stained a pinkish 
hue. The adult male taken in June is in fresh plumage ; but the young male, also 
taken in June, is in full moult. In this bird the new feathers of the under-parts, unlike 
the worn ones, are strongly barred like those of the adults. The adult male from the 
Hagen Range is topotypical and both it and the other adult male show the character- 
istics of captus given by Mayr and Gilliard, but the wings average a little longer 
(wing, <$ 95-102), like those of specimens taken at Nondugl by Gyldenstolpe (1955 : 
32). The adult female, in comparison with the adult male, has the back paler in 
colour by being a clearer olive-brown, less reddish. In the juvenile male the crown 
and the neck lack the maroon-chestnut of the adult and there is a reduction in the 
white barring of the back. In this respect the young bird agrees with a series of 
adults of R. p. pectoralis (Temminck) from eastern Australia. 

Rallus philippensis wahgiensis Mayr and Gilliard. 

Rallus philippensis wahgiensis, Mayr and Gilliard, 1951, Amer. Mus. Novit. No. 1524, p. 3. 
Nondugl, Central Highlands, New Guinea. 

1953.17.46 ; i <$ ; Mount Giluwe ; 7,500 ft.; July, 195 1. 

Measurements : 1 $ — wing, 153 ; tail, 76 ; bill, 31 ; tarsus, 41. Colours : bill, 
dark horn, base of lower mandible reddish ; legs, light brown ; iris, reddish brown. 
Stomach contents : insects. 

This specimen differs only in size from a topotypical male collected by Mr. Gilliard. 
The wing and the tail are longer and the tarsus shorter, the measurements of 


wahgiensis given by Mayr and Gilliard are : wing, 137, 140 ; tail, 62.5 ; tarsus, 44, 
45. In this respect it matches specimens taken at Nondugl by Gyldenstolpe (1955 : 32) . 

Porzana pusilla palustris Gould 

Porzana palustris Gould, 1843, Proc. Zool. Soc. Lond. 1842, p. 139. Tasmania. 

Porzana pusilla mayri Junge, 1953, Zool. Meded., Leiden, 31, p. 247. Paniai, western New 

1953.17.47-49 ; 2 ?, i juv. $ ; Mount Giluwe, 8,000-7,500 ft.; May, June, 1951. 

Measurements : 3 $ — -wing, 79-83 ; tail, 35-40 ; bill, 14-3-15. Colours : bill, 
olive-green, tip horn colour ; legs, olive-green ; iris, orange-red. Stomach contents : 
small insects. 

The Mount Giluwe birds may be distinguished from a series of palustris from 
eastern Australia on very slight colour differences. The upper-parts of the New 
Guinea birds are somewhat warmer in tone and the under-parts a slightly clearer 
grey, but these differences may be attributable to the freshness of the skins. Junge 
(1952 : 247) was the first to report the presence of this species in New Guinea. He 
distinguished a new race, mayri, on a series from Paniai separating it from the Austra- 
lian birds on a greater wing length. The measurements of the material before me, 
however, do not support this separation for there is some variation. 

Australian birds, in B.M. (N.H.) 

„ (Junge, 1952 : 247) 

New Guinea ,, in B.M. (N.H.) 

„ „ (Junge, 1952 : 247) 

Wing, 75-81 1 

„ 80-85/ 75 ° 5 

" T 83 >76-8 3 
„ 76-79/ / J 

Porzana tabuensis edwardi Gyldenstolpe 

Porzana tabuensis edwardi Gyldenstolpe, 1955, Ark. f. Zool., Ser. 2, 8, p. 3. Nondugl, Central 
Highlands, New Guinea. 

1953.17.50-56; 2 (J, i juv. (J ; 2 ?, 2 juv. ?; Mount Giluwe, 8,000-7,5000 ft.; 
March, April, June, 195 1. 

Measurements : 3 <$ — wing, 78-85 ; tail, 41-50 ; bill, 18-19. 4 ¥ — wing, 
78-80 ; tail, 41-46 ; bill, 16-18. Colours : bill, black ; legs, coral-red ; iris, orange- 
red. Stomach contents : insects and grass seeds. 

Gyldenstolpe (1955 : 34) separated the birds of the Wahgi Valley on their darker 
colour and slightly longer bill in comparison with a series of tabuensis from the Tuamoto 
Archipelago and Rapa Island. These characters are present in the Mount Giluwe 
birds and distinguish them from a series from various Pacific islands. The adult male 
taken in April is in worn plumage, while that taken in June is in fresh. The fresh 
plumage is more vinous above and more blue than slate-grey below. The young male, 
31st March, differs from the adult in having the chin and throat white and the 


abdomen and under- wing coverts a pale whitish-grey. This bird is in moult. The adult 
females taken in April are starting moult with a few new feathers coming through on 
the back. Of the two young females one, 5th April, has nearly completed moult ; 
while the other, 7th April, has only a few new feathers coming through on the back. 

Rallicula forbesi subspecies? 

Rallicula forbesi Sharpe, 1887, in Gould's "Birds of New Guinea", pt. 23, pi. (70). Owen Stanley 
Range, south-western New Guinea. 

1953.17.57 ; i(J; Hagen Range ; 8,000 ft.; February, 1951. 

1953 . 17 . 58-62 ; 1 (J, 4 $ ; Mount Giluwe ; 7,400-7,300 ft.; May, June, July, 1951. 

Measurements : 2 $ — wing, 111-112 ; tail, 77-79 ; bill, 24-25. 4 $ — wing, 
108-112 ; tail, 63-68 ; bill, 24-26. Colours : bill, brown-black ; legs, brown-black 
to black ; iris, pale to dark brown. 

The two males differ only very slightly from the type and a series of forbesi from 
south-eastern New Guinea in that the plumage is somewhat richer in tone. The four 
females, on the other hand, exhibit an interesting variation. Although they appear to 
be inseparable from the allotype and a series of forbesi in most respects there are 
differences in the speckling and the occurrence of a red-chestnut fringe on the feathers 
of the back. The July bird agrees with the allotype in colour and pattern in that over 
the back the speckling is the typical buff colour and many of the feathers are fringed 
imperfectly with a red-chestnut colour. Gradations are found in the other females to 
the colour and pattern of a May bird (1953. 17.62) in which the spotting is no longer 
buff but white, and replaced over the scapulars by white streaks ; also none of the 
feathers of the back have a red-chestnut fringe. The markings of this specimen so 
approached the description of the race steini which Rothschild described on one female 
from the Weyland Mountains that it was sent to the American Museum to be compared 
with the unique type. There Dr. Amadon, in Mr. Gilliard's absence, kindly examined 
the specimens and found that the Mount Giluwe bird matched the type. Mayr and 
Gilliard (1954 : 335) described a similar variation in a Wahgi region series which they 
named steini mainly on the length of the tail. The present series, however, cannot be 
separated from the nominate form on measurement, while on plumage most of the 
birds approach forbesi. It seems that if the present series were combined with the 
birds reported by Mayr and Gilliard then phenotypical forbesi and steini would be 
present in approximately equal numbers in addition to the specimens grading 
between the two. In view of this I do not venture to name the birds of the present 
series at racial level. 

One explanation of the occurrence of this variation in these series from the Wahgi 
region could be that the two races have come into breeding contact following a period 
of reproductive isolation. It seems that such an interpretation, based on the material 
available, would not be entirely incorrect because a population showing such a high 
incidence of heterogeneity would be unlikely to occur in a cline. It seems equally 


unlikely that the type of steini and other specimens that resemble it are variants for 
no similarly patterned birds have been reported from southeast of the Wahgi region. 

Pluvialis dominca fulva (Gmelin) 

Charadrius fulvus Gmelin, 1789, Syst. Nat. 1, pt. 2, p. 687. Tahiti. 

Field Note. Six of these plovers were seen together in a " paddock " at Nondugl 
on 15th September. A few of the black feathers of the breeding plumage could be 
seen on the under-surface of some of the birds. 

Scolopax saturata rosenbergii Schlegel 

Scolopax rosenbergii Schlegel, 1871, Ned. Tijdschr. Dierk. 4, p. 54. Arfak Mts., north-western 
New Guinea. 

1953.17.63-65; 2 <J, 1 ? ; Mount Giluwe, 9,500-9,000 ft.; May, June, 1951. 

1953.17.66-67 ; 1 (J, 1 ? ; Lamende Range, 9,500-9,000 ft.; June, 1951. 

Measurements : 2 <J — wing, 145-146 ; tarsus, 33-34 ; bill, 82-83. 3 $ — wing, 
148-155 ; tarsus, 32-35 ; bill, 83-87. Colours : bill, brown-black horn ; legs, 
dark brown-grey ; iris, dark brown. Stomach contents : worms, grubs, small 
beetles and other insects. 

" The Woodcock is a bird of the high mountain-forest on Mount Giluwe and the 
Lamende Range. It lives on the ground, where it searches for insects and digs for 
worms and grubs. During the early morning it will often rise and fly high above the 
trees, where it can be heard calling ; later it drops down again on to the forest floor ' ' 
(F. S.-M.). 

The black barring of the feathers of the back is appreciably heavier than in topo- 
typical specimens of rosenbergii. In this respect the present series is similar to the 
specimens that I have seen from the Oranje Mountains taken by Dr. Rand who 
considered that this difference was of no racial significance. (19426 : 439). The wing 
measurements are less that those of Dr. Rand's specimens but Shaw-Mayer's birds are 
in moult and it seems that the primaries are not fully grown. In these moulting birds 
the brown of the new feathers is noticeably redder and darker than that of the old. 

Gallicolumba beccarii beccarii (Salvadori). 

Chalcophaps beccarii Salvadori, 1875, Ann. Mus. Civ. Genova, 7, p. 974. Hatam, Arfak Moun- 
tains, north-western New Guinea. 

1953.17.68 ; i (J ; Hagen Range, 8,000 ft.; March, 1951. 

1953.17.69 ; 1 c? ; Mount Giluwe, 7,500 ft.; May, 1951. 

Measurements : 2 ad. <J — wing, 107,109 ; tail, 60,70 ; bill, 14. Colours : bill, 
black ; legs, dark purple ; iris, brown-black. Stomach contents : hard seeds. 


Charmosyna papou goliathina Rothschild and Hartert 

Chavmosyna stellae goliathina Rothschild and Harter, 191 1, Novit. Zool. 18 ,p. 160. Mt. Goliath, 
Oranje Range, central New Guinea. 

1953 . 17 . 70-73 ; 2 (J, 2 $ ; Mount Giluwe, 9,000-8,500 ft.; April, June, July, 1951. 

Measurements : 2 (J— wing, 151, 155 ; tail, 297, 324 ; bill, 18. 2 ? — wing, 150, 
152 ; tail, 256, 280 ; bill, 17, 19. Colours : bill, orange to dark red ; legs, bright 
orange ; iris, orange-red. Stomach contents : flowers and fruit pulp. 

The two males are in the dark phase plumage. The June and July birds of both 
sexes are completing moult. 

Field note. Native name, " Kai-niginch ". 

Oreopsittacus arfaki grandis Ogilvie-Grant 

Oreopsittacus grandis Ogilvie-Grant, 1895, Bull. Brit. Orn. Club, 5, p. 15. Owen Stanley 
Mountains, south-eastern New Guinea. 

1953.17.74-76 ; i <J, 2 ? ; Hagen Range, 9,500-9,000 ft.; December, 1950. 

Measurements : 1 <J — wing, 81 ; tail, 84 ; bill, 8. 2 $ — wing, 82, 86 ; tail, 85 ; 
bill, 8, 9. Colours : bill, black ; legs, green-grey ; iris, black-brown. 

These birds match grandis except in wing length. In this respect the females are 
larger than both grandis and major and the males intermediate, — grandis : $, 73-76 ; 
9, 73 — major : (J, 85-87 ; $, 80 — (Ogilvie-Grant, 1915 : 169). 

Neopsittacus musschenbroekii major Neumann 

Neopsittacus musschenbroekii major Neumann, 1924, Ornith. Monatsber. 32, p. 38. Schrader- 
berg, Sepik Mountains, northern New Guinea. 

I953-I7-77 ; 1 (J ; Mount Giluwe, 7,500 ft.; June, 1951. 

Measurements : 1 <$ — wing, 114 ; tail, 101 ; bill, 16. Colours : bill, yellow ; 
legs, brown-grey ; iris, orange-red. Stomach contents : fruits and berries. 

Other birds taken in the same area were identified as major by Mayr and Gilliard 
(1954 : 339) but the present example differs from other material of the species in 
being much darker. Compared with major, the crown, collar, nape and sides of the 
throat are browner and washed with crimson. The chin and throat are dark brown 
with a greenish wash. The back appears to be very dark due to the feathers being 
fringed with dark brown. Some of the primary wing coverts, the outer webs of the 
primaries and the upper surfaces of the two central rectrices are almost entirely dark 
brown with a slight greenish wash. The pattern of the underparts is similar to major, 
but the green is replaced by olive and the crimson by a yellowish-brown. This 
specimen may be an abnormally pigmented bird of the race major, although the 
possibility that it represents a distinct form should not be overlooked. 


Psittacella brehmii pallida A. B. Meyer 

Psittacella pallida A. B. Meyer, 1886, Zeitschr. ges. Ornith. 3, p. 3. Astrolabe Mountains, 
south-eastern New Guinea. 

Psittacella biirgersi Reichenow, 1918, Journ. f. Ornith. 66, p. 244. Schraderberg, Sepik region, 
northern New Guinea. 

Psittacella brehmi ornata Mayr, 193 1, Mitt. Zool. Mus. Berlin, 17, p. 720. Dawong, Herzog 
Mountains, south-eastern New Guinea. 

1953.17.78-79 ; i <J, i $ ; Mount Giluwe, 8,500 ft.; June, 1951. 

1953. 17.80-81 ; 1 (J, 1 juv. $ ; Hagen Range, 9,000 ft.; February, 1951. 

Measurements : 2 <$ — wing, 122, 125 ; tail, 86 ; bill, 18, 20. 1 ? — wing, 127 ; 
tail, 87 ; bill, 18. 1 juv.? — wing, 122 ; tail, 86 ; bill, 17. Colours : bill, metallic 
blue-grey, becoming white towards tip in juvenile ; legs, dark grey, green-grey in the 
ad. (J ; iris, orange, grey in juvenile. Stomach contents : hard berries and seeds. 

Gyldenstolpe (1955 : 57) described much of the individual variation in both color- 
ation and size in the birds of this species from central and south-eastern New Guinea. 
He showed that biirgersi should be regarded as a synonym of pallida and agreed with 
Mayr (1941 : 69) who had previously united ornata with pallida. Of the present 
specimens the adult female and the juvenile were sent to the Berlin Museum, where 
Professor Stresemann, who kindly made the comparison, found that they closely 
matched the type of P. biirgersi Reichenow. In view of the evidence presented by 
Gyldenstolpe I am naming the birds of this present series pallida. Compared with a 
small series of intermedia from southern Dutch New Guinea the Wahgi region birds 
are more bluish-green above with a greenish wash over the crown ; and the sides of 
the head are more greyish in colour. 

Psittacella picta excelsa Mayr and Gilliard 

Psittacella picta excelsa Mayr and Gilliard, 1951, Amer. Mus. Novit. No. 1524, p. 6. Mount 
Orata, Kubor Mountains. Central Highlands, New Guinea. 

1953.17.82 ; I (J ; Hagen Range, 9,000 ft.; February, 1951. 

1953.17.83 ; 1 juv. $ ; Kubor Range, 8,000 ft.; October 1950. 

1953.17.84 ; 1 (J ; Mount Giluwe, 11,000 ft.; April, 1951. 
Measurements : 2 <J — wing, 114, 116 ; tail, 73 ; bill, 14, 15. 1 juv. $ — wing, 

103 ; tail, 68 ; bill, 12-5. Colours : bill, pale blue, yellowish (juv. (J) ; legs, grey ; 
iris, orange-yellow, grey-yellow (yg. cJ). Stomach contents : seeds and small hard 

The adult from the Hagen range differs from the Mount Giluwe bird and a topo- 
typical specimen taken by Mr. Gilliard in that the golden-yellow patches on each 
side of the neck are joined over the neck by a narrow band of the same colour. The 
young male resembles a topotypical female also collected by Mr. Gilliard. 

ZOOL. 3, IO. 26 



Psittacella modest a hallstromi Mayr and Gilliard 

Psittacella modesta hallstromi Mayr and Gilliard, 1951, Amer. Mus. Novit. No. 
Yandarra, north slope of Mount Wilhelm, Bismarck Mountains, New Guinea. 


P- 5 

1953.17.85-87 ; 2 (J, 1 $ ; Mount Giluwe, 8,500-8,000 ft.; April, June, 1951. 

1953.17.88 ; 1 juv. $ ; Rubor Range, 8,000 ft.; October, 1950. 

1953.17.89 ; 1 <J ; Hagen Range, 9,500 ft.; February, 1951. 
Measurements : 4 <J — wing, 98-101 ; tail, 59-62 ; bill, 14. 1 ? — wing, 102 ; 

tail, 64 ; bill, damaged. Colours : bill, metallic blue-grey ; legs, dark blue-grey ; 
iris, orange (2 ad. (J), yellow-brown (2 imm. (J), dark brown (1 ad. ?). Stomach 
contents : seeds and pieces of hard berries. 

The young male taken in October is starting to moult from juvenile plumage in 
which the throat and breast are an olivaceous colour and lack the ashy-brown of the 
adult male. The feathers of the breast are also faintly barred with yellow as in the 
female where the orange and black barring of the breast merges into the plain green 
of the abdomen and flanks. The head and the nape are the same colour as the back 
although the feathers at the base of the bill are washed with blue and some on the 
hind neck are heavily barred with orange and black. The February bird is in worn 
plumage, while the April and June birds are in fresh plumage. The testes of the June 
male are enlarged. 

Caco mantis pyrrhophanus excitus Rothschild and Hartert 

Cacomantis excitus Rothschild and Hartert, 1907, Novit. Zool. 14, p. 436. Angabunga River, 
south-eastern New Guinea. 

1953. 17.90-91 ; i (J, i juv. (J ; Mount Giluwe, 7,500 ft.; April, May, 1951. 

Measurements : 1 $ — wing, 145 ; tail, 140 ; bill, 17. 1 juv. <J — wing, 122 ; tail, 
100 ; bill, 14. Colours : bill, black ; legs, yellow ; iris, dark brown. Stomach 
contents : caterpillars and flying insects. 

These birds were taken in the wooded country on the north slopes of Mount Giluwe. 
The May bird is similar to a male from the Utakwa River in Dutch New Guinea 
except that the grey of the chin extends to the throat. The young male which is in 
fresh plumage (April) is plain chestnut above and rufous-brown irregularly barred 
with black below. 

Chalcites meyerii (Salvadori) 

Chrysococcyx meyerii Salvadori, 1874, Ann. Mus. Civ. Genova, 6, p. 82. New name for Chryso- 
coccyx splendidus A. B. Meyer, 1874, Sitzungsber, Akad. Wiss. Wien, 69, p. 81. Hatam, 
Arfak Mountains. (Not Gray, 1847). 

1953.17.92 ; 1 c? ; Wahgi River ; 5,000 ft.; November, 1950. 
Measurements : 1 ? — wing, 88 ; tail, 65 ; bill, 12-5. Colours 
legs, blue-grey ; iris, dark brown. 

bill, black 


Tyto tenebricosa arfaki (Schlegel) 

Strix tenebricosa arfaki Schlegel, 1879, Notes Leyden Mus. 1, p. 101. Arfak Mountains, north- 
western New Guinea. 

1953.17.95 ; i (J ; Hagen Range, 8,500 ft.; March, 1951. 

1953.17.96 ; 1 ? ; Lamende Range, 8,500 ft.; March, 1951. 
Measurements : 1 $ — wing, 275 ; tail, 119 ; tarsus, 62. 1 ? — wing, 292 ; tail, 

130 ; tarsus, 73. Colours : bill, dark horn (ad. $, lower mandible pale horn) ; legs, 
dark flesh ; iris, brown-black. Stomach contents : 1 (J, remains of a rat. 

Tyto longimembris papuensis Hartert 

Tyto longimembris papuensis Hartert, 1929, Novit. Zool. 35, p. 103. Owgarra, Angubunga 
River, south-eastern New Guinea. 

1953.17.93-94 ; 1 <J, 1 $ ; Mount Giluwe, 7,400 ft.; May, 1951. 

Measurements : 1 ad. $ — wing, 343 ; tail, 114 ; tarsus, 84. 1 ad. ? — wing, 357 ; 
tail, 123 ; tarsus, 97. Colours : not recorded. Stomach contents : nil. 

" <5, taken from nest in tree-hollow brooding three well incubated eggs ; time, 
10 a.m. $, taken near hollow containing nest ; time, 10 a.m. Usually one or two of 
these Grass Owls could be seen flying low over the grasslands or sitting on an old tree 
fern stump during the late afternoon or at dusk. The species was, however, uncommon 
on the plateau, possibly the altitude was a little too great for it. It was plentiful on 
the grasslands east of the Hagen Range at about 5,000 feet." (F. S.-M.). 

It is interesting that the male was taken while brooding the eggs as it is generally 
accepted that it is not usual for the males of this species to share in incubation. The 
eggs were " well incubated " on the 26th May, so it appears that the breeding season 
of the New Guinea bird is similar to that of walleri, from Australia, which breeds 
during May and June. It is worth mentioning, perhaps, that in November, 1946, Mr. 
Shaw-Mayer collected a pair of Grass Owls at Yanka and in comparison the birds 
from Mount Giluwe are slightly more tawny above and none of the feathers of the 
back have white shaft streaks. The male from Yanka resembles the females in that 
the feathers of the sides of the face-ring and the chin are tipped with brown and the 
face is washed with the same colour, whereas these parts are white in the male 
from Mount Giluwe. 

Ninox theomarcha theomarcha (Bonaparte) 

Spiloglaux theomarcha Bonaparte, 1885, Compt. Rend. Acad. Sci. Paris, 41, p. 654. Triton 
Bay, south-western New Guinea. 

1953.17.97 ; 1 juv. $ ; Hagen Range, 8,200 ft.; February, 1951. 
Measurements : 1 juv. $ — wing, 188 ; tail, 98 ; tarsus, 29. Colours : bill, dark 

horn ; legs, pale green-flesh ; iris, bright yellow. Stomach contents : remains of 
large insects. 


The bird has started post-juvenile moult with new feathers coming through on the 
breast, but towards the vent the paler juvenile plumage persists. 

Podargus papuensis Quoy and Gaimard 

Podargus papuensis Quoy and Gaimard, 1830, Voy. Astrolabe, Zool. 1, p. 207 (pi. 13). Dorey, 
Vogelkop, north-western New Guinea. 

1953.17.98 ; 1 $ ; Mount Giluwe, 7,300 ft.; May, 1951. 

Measurements : 1 ad. ? — wing, 285 ; tail, 271 ; tarsus, 20. Colours : bill, 
horn ; legs, dark grey ; iris, dark orange-red. Stomach contents : beetles. 
Taken in the woodlands at the north base of Mount Giluwe. 

Aegotheles albertisi salvadori Hartert 

Aegotheles salvadori Hartert, 1892, Cat. Bds. Brit. Mus. 16, p. 649. Astrolabe Mountains, 
south-eastern New Guinea. 

1953 . 17 . 99-103 ; 2 (J, i $, 2 juv. ? ; Hagen Range, 8,500 ft.; February, 1951. 

1953 . 17 . 104-105 ; 1 juv. (J, 1 juv. $ ; Mount Giluwe, 7,500 ft.; April, May, 1951. 

Measurements : 3 <$ — wing, 118-121 ; tail, 97-98 ; bill, 8-9. 4 $ — wing, 
116-125 ; tail, 97-102 ; bill, 8-9. Colours : bill, upper mandible dark horn and 
lower mandible pale horn ; legs, dark flesh ; iris, dark brown. Stomach contents : 

Nearest to salvadori but like the specimens described by Mayr and Gilliard 
(1954 : 342) they differ from the typical form by the underparts being somewhat more 
coarsely marked. The younger birds are moulting, the older are mostly in worn 

Aegotheles insignis insignis Salvadori 

Aegotheles insignis Salvadori, 1875, Ann. Mus. Civ. Genova, 7, p. 916. Hatam, Arfak Mountains, 
north-western New Guinea. 

1953. 17. 106 ; i c? ; Mount Wilhelm, Bismark Range ; 6,000 ft.; May, 1951. 
1953.17.107-111 ; 2 <J, 1 juv. cJ, 3 $ ; Hagen Range, 8,500-8,000 ft.; February, 
March, 195 1. 

1953.17.112-114 ; 1 juv. (J, 2 ? ; Mount Giluwe, 8,000-7,500 ft.; March, June, 


Measurements : 4 <$ — wing, 157-170 ; tail, 132-142 ; tarsus, 21-22. 5 °. — wing, 
158-177 ; tail, 135-155 ; tarsus, 21-23. Colours : bill, upper mandible dark 
brown and lower mandible pale horn ; legs, flesh ; iris, dull yellow to brown. 
Stomach contents : beetles and other insects. 


This series exhibits the colour phases of this race reported by Hartert (1930 : 95) 
and Rand (19426 : 456). There are three light phase and six dark phase birds, three 
of the latter being considerably darker than any I have seen before. 

Eurostopodus archboldi (Mayr and Rand) 

Lycornis archboldi Mayr and Rand, 1935, Amer. Mus. Novit. No. 814. p. 4. Mount Tafa, 
Wharton Range, south-eastern New Guinea. 

1953. 17. 119 ; i ^ ; Tomba, Hagen Range, 8,000 ft.; February 1951. 

1953.17.120-121 ; 2 $ ; Mount Giluwe, 7,500-7,300 ft.; May, June, 1951. 

Measurements : 1 $ — wing, 120 ; tail, 156 ; bill, 9-5. 2 $ — wing, 123, 125 ; 
tail, 145, 149 ; bill, 10. Colours : bill, black ; legs, dark grey ; iris, brown-black. 
Stomach contents : insects. 

The male is in worn plumage and there is little spotting on the breast because many 
of the feathers have lost their rufous tips. The May female is in fresh plumage and 
shows numerous breast spots. A further difference between these specimens seems 
to be due to fading for the colour of these spots is paler in the male than in the female. 
These differences are interesting because they are qualitatively the same but quanti- 
tatively greater than those between two specimens collected by Dr. Rand ; namely, 
a male taken at Lake Habbema in Dutch New Guinea and a topotypical female 
taken in September. These match the male and the May female respectively of the 
present series, even in the plumage of the male being more worn than that of the 
female. Rand (19426 : 457) suggested that the differences between these specimens 
might indicate that the Dutch New Guinea birds form another race, but on this point 
he was not certain. The evidence provided by the birds from the Central Highlands 
suggests that these are seasonal differences. Dr. Rand also noted a slight difference 
in the colour and clarity of the black markings and vermiculations of the scapulars 
and inner secondaries of his birds. In all the birds of the present series these tend to 
be closer to the Lake Habbema bird in colour and to the Mount Tafa bird in clarity, 
so it may well be that this small sample is from a mixed population. 

Collocalia esculenta esculenta (Linnaeus) 

Hirundo esculenta Linnaeus, 1758, Syst. Nat. 10th ed., p. 191. " China " error for Ambon 
{ex Rumphius). 

Field notes : These swiftlets often occurred in pairs ; once, however, about 12 
were seen hawking for insects above a native clearing on a steep hillside at about 6,000 
ft.. A nest was found under a bank beside a stream, in it were two elongate eggs a 
pale blue-white in colour (10th September). The nest was made of fine roots and 
thickly covered with a yellowish coloured dry lichen that hung from the trees. 


Collacalia hirundinacea hirundinacea Stresemann 

Collocalia fuciphaga hirundinacea Stresemann, 1914, Verh. Ornith. Ges. Bayern, 12, p. 7. 
Setakwa River, Dutch New Guinea. 

Field note : A few birds were seen, two being found on the nest. The nests were 
built of fine rootlets, thin black stalks and a few pieces of moss with some earth 
towards the bottom. An elongate egg was found in one of the nests. 

Alcyone azurea ochrogaster Reichenow 

Alcyone ochrogaster Reichenow, 1903, Journ. f. Ornith. 51, p. 149. Ramu River, northern New 

1953. 17. 115 ; i ? ; Minj, Wahgi River, 5,000 ft.; November, 1950. 
Measurements : 1 ? — wing, 77 ; tail, 33 ; bill, 46. Colours : bill, black ; tip 

pale horn ; legs, orange-red ; iris, not recorded. Stomach contents : insects. 

This bird matches ochrogaster except that the upper-parts are a duller blue like 

Clytoceyx rex rex Sharpe. 
Clytoceyx rex Sharpe, 1880, Ann. Mag. Nat. Hist. (5) 6, p. 231. East Cape, New Guinea. 

1953. 17. 116 ; i ? ; Mount Wilhelm, Bismarck Range, 6,000 ft.; May, 1950. 
Measurements : 1 $ — wing, 173 ; tail, 137 ; bill, 37. Colours : bill, pale horn 

(base of upper mandible dark horn) ; legs, green-grey ; iris, black-brown. Stomach 
contents : remains of large beetles. 

This bird is more richly coloured than others I have seen of the species. This is 
interesting because Paludan described septentrionalis on the paler undersurface of 
three specimens from the nearby Sepik area. In size it is intermediate between rex 
and imperator van Oort from the Noord River — imperator,9. : wing, 180 ; tail, 148 ; 
(van Oort, 1909 : 79) — rex, $ : wing, 160-165 ; tail, 115-125. 

Halcyon sancta sancta Vigors and Horsfield. 

Halcyon sancta Vigors and Horsfield, 1827, Trans. Linn. Soc. London, 15 ; p. 206. New South 
Wales, Australia. 

1953.17.117-118 ; 2 (J ; Mount Giluwe, 7,600-7,500 ft.; May, 1951. 

Measurements : 2 (J— wing, 91, 92 ; tail, 59, 60 ; bill, 35, 37. Colours : bill, 
black (lower mandible pale towards base) ; legs, dark-grey ; iris, dark-brown. 

These non-breeding visitors were taken in the wooded hills north of Mount Giluwe. 
Both are in moult, 


Hirundo tahitica frontalis Quoy and Gaimard 

Hirundo frontalis Quoy and Gaimard, 1830, Voy. Astrolabe, Zool. 1, p. 204, pi. 12. Dorey, 
north-western New Guinea. 

Field Note. About 15 to 20 of these birds were to be seen on most evenings on 
the earthy bank around the pond at Nondugl (8th September) . 

Edolisima montona montona (Meyer) 

Campephaga montona A. B. Meyer, 1874, Sitzungsber. Akad. Wiss, Wein, 69. p. 386. Arfak 
Mountains, north-western New Guinea. 

1953. 17. 122 ; 1 c? ; Hagen Range, 8,000 ft.; March, 1951. 

Measurements : 1 <$ — wing, 135-5 \ tail, 101 ; bill, 16. Colours : bill, black ; 
legs, black ; iris, black-brown. Stomach contents : berries. 

Rothschild and Hartert (1907 : 464) examined birds of this species and found that 
the adult males from the mountains of western New Guinea had longer wings than 
those from the mountains of the south-east ; they named these shorter-winged birds 
minus. This difference in wing length, however, does not appear to be so well marked 
now that more material is available ; Junge (1953 : 40), for example, reported that 
the wing lengths of some adult males in a series from one locality in western New 
Guinea were within the range of measurements of minus. Nevertheless, until more 
material is available from other localities the species cannot be reviewed satisfactorily 
but it seems probable that when this is done minus will be found to be no longer a 
valid race separable on wing length. The specimen from the Hagen Range is similar 
in measurement to some of the birds reported by Junge. 

Wing Length of Adult Males 

Western New Guinea. South-eastern New Guinea. 

Rothschild and Hartert . 136-5-141 -5 Rothschild and Hartert . . 128-132 

Junge .... 131 -141-5 British Museum (Natural History) 125-134 

Coracina longicauda longicauda (De Vis) 

Grauculus longicauda De Vis, 1890, Ann. Rep. Brit. New Guinea, 1888-89, p. 59. Musgrave 
Range, south-eastern New Guinea. 

Field note. A nest was found about 50-60 ft. above the ground in a fork of a 
horizontal branch on a somewhat isolated tree in the forest near Tomba. It was 
saucer-shaped and built of fine rootlets, the outside was decorated with lichens and 
moss. There was one juvenile on n October, which was fed by both parents. The 
food appeared to consist mainly of winged insects, but on two occasions (13th October) 
lizards were brought. After being fed with the first one, which had been pushed down 
its throat with some difficulty, the young bird made a gurgling noise which may have 


been a begging call somewhat stifled by the full crop. On the second occasion when a 
lizard was brought to the nest only the tail protruded from the adult's mouth, the 
adult then regurgitated the lizard to give it to the young. When the nest was examined 
on the following day (14th October) an uneaten green lizard was found there. The 
nest was kept under observation for about three and a half hours and the feeding 
visits that the adults paid to the nest during that time were recorded as follows : 


Male and female. 

10.30 : 




10.42 : 




11.25 : 

Male and female 


Male and Female. 

12.07 : 


Nest sanitation was attempted only once, but that may have been because the 
birds had been disturbed. When it was not being fed the young bird remained 
wonderfully still, it never changed its position during the whole of the morning while 
it was being watched from the hide. On another day the bird was moved around in 
its nest so that it faced the sun, it turned immediately and resumed its former 

Anthus gutturalis rhododendri Mayr 

Anthus gutturalis rhododendri Mayr, 1931, Mitt. Zool. Mus. Berlin, 17, p. 692. Mongi-Busu, 
Saru waged Mountains (2,600 m.), north-eastern New Guinea. 

1953. 17. 123-124 ; i (J, i juv. (J ; Hagen Range, 11,000 ft.; February, 1951. 

1953.17.125-131 ; 4 c?, 3 ? ; Mount Giluwe, 12,000-11,500 ft.; April, 1951. 

Measurements : 5 <$ l — wing, 98-101 ; tail, 74-79 ; bill, 12-14. 3 $ — wing, 
94-97 ; tail, 74-77 ; bill, 13-14. Colours : bill, black ; legs, dark horn ; iris, 
dark brown. Stomach contents : grass seeds and insects. 

1 Measurements of 1953. 17. 124 not given owing to broken plumage. 

Anthus australis exiguus Greenway 

Anthus australis exiguus Greenway, 1935, Proc. New England Zool. Club, 14, p. 35. Wau, 
Morobe district, north-eastern New Guinea. 

1953. 17. 132-138 ; 3 <J, 3 $, 1 juv. ? ; Mount Giluwe, 7,300 ft.; May, June, 1951. 

Measurements : 3 $ — wing, 84-86 ; tail, 62 ; tarsus, 26-27 ; pollux (not claw), 
9-9-5 ; claw of pollux, 10-13 \ bill, 13-5-14. 3 ? — wing, 80-82 ; tail, 55-59 ; tarsus, 
26 ; pollux (not claw), 9 ; claw of pollux, 11-13 ; bill, 12-14. 1 juv $ — wing, 76 ; 
tail, 53 ; tarsus, 25 ; pollux (not claw), 9 ; claw of pollux, 12 ; bill, 13. Colours : 
bill, dark horn (lower mandible paler towards base) ; legs, dark flesh to horn ; iris, 
dark brown. Stomach contents : insects. 

" This pipit was plentiful on the plateau at just over 7,000 feet. It was especially 
numerous on any cleared area of grassland and in the native gardens. As many as 
50 to 60 were observed in the early morning and late afternoon on a recently made 
football field near lumbal." (F. S.-M.). 


No colour difference can be seen between most of the males and females of this 
series and the type and allotype respectively of exiguus kindly sent on loan to me by 
Mr. Greenway, One adult male, however (1953 . 17 . 134) is slightly paler, but warmer 
in colour above owing to the light brown edging to the feathers being wider than in the 
type ; the breast and flanks are correspondingly warmer and are washed a light 
brown. They are also slightly larger, particularly in the length of the wing and tail, 
but probably not enough to be recognized racially. 

Field note. During September they were seen in pairs on the lawn around the 
house at Nondugl and on the grasslands where they were very common. One bird 
was seen in partial display, it flew typically pipit-like into the sky and then volplaned 

Saxicola caprata wahgiensis Mayr and Gilliard 

Saxicola caprata wahgiensis Mayr and Gilliard, 1951, Amer. Mus. Novit. No. 1524, p. 8. 
Mafulu, south-eastern New Guinea. 

1953. 17. 139-142 ; 2 (J, 2 $ ; Mount Giluwe, 7,300 ft.; May, June, 1951. 

Measurements : 2 <J — wing, 77, 79 ; tail, 55, 56 ; bill, 11. 2 $ — wing, 77, 78 ; 
tail, 56, 58 ; bill, 11. Colours : bill, black ; legs, black ; iris, brown-black. 
Stomach contents : insects (grubs and adults). 

" Two or three of these friendly stone-chats are usually to be seen around any 
building in the grasslands on the plateau. Their song is the first bird call to be heard 
in the early morning." (F. S.-M.). 

Compared with the female specimen of wahgiensis collected by Mr. Gilliard at 
Nondugl the two females from Mount Giluwe are slightly darker below and on the 
sides of the neck. In the darker colour of the underparts these birds tend towards the 
Oranje Mountain bird, belensis, of which the females are larger than the present pair, 
Mayr and Gilliard (195 1 : 8). 

Field note. Native name, " Kompichente ". A nest was found built on the 
ground on a pile of pit-pit roots and stems. It contained two juveniles with the 
feathers of the first plumage newly broken through the skin (26th September) . Only 
the female was seen to visit the nest. A pair was seen in the garden perched on 
sticks in the flower beds, each dropped down tc the ground from time to time to pick 
up insects. A fairly tame bird often seen in the Mount Hagen area. 

T urdus poliocephalus erebus Mayr and Gilliard 

Turdus poliocephalus erebus Mayr and Gilliard, 1952, Amer. Mus. Novit. No. 1577, p. 7. New 
name for Turdus poliocephalus carbonarius Mayr and Gilliard, 195 1, Amer. Mus. Novit. No. 
1524, p. 7. Mount Orata, Kubor Mountains, Central Highlands, New Guinea. (Not 
Turdus carbonarius Lichtenstein, 1823.) 

1953 . 17 . 143-146 ; 3 (J, 1 $ ; Mount Giluwe, 11,000 ft.; April, 195 1. 

Measurements : 3 $ — wing, 124-132 ; tail, 90-94 ; bill, 20-21. 1 ? — wing, 123 ; 
tail, 92 ; bill, 21. Colours : $ — bill, orange-yellow with darker tips ; legs, orange ; 
iris, brown with yellow eye ring ; inside of mouth, orange. ? — bill, upper mandible 


dark horn, lower mandible orange ; legs, dull orange ; iris, dark brown — yellow 
eye ring. Stomach contents : berries and small beetles. 

" On the top of Mount Giluwe the blackbird leaves the shelter of the small timber 
in the late afternoon and flies into the open alpine grassland where it searches among 
the rocks and tussock-grasses for its food." (F. S.-M.). The gonads of all the 
specimens were enlarged. 

Melampitta lugubris longicauda Mayr and Gilliard 

Melampitta lugubris longicauda Mayr and Gilliard, 1952, Amer. Mus. Novit. No. 1577, p. 1. 
Mount Tafa, Wharton Range, south-eastern New Guinea. 

1953 . 17 . 147-148 ; i juv. (J, i ? ; Mount Giluwe, 9,000-8,500 ft.; April, 1951. 

Measurements : 1 juv. <$ — wing, 83 ; tail, 33 ; tarsus, 40 ; bill, 15. 1 
§ — wing, 84 ; tail, 54 ; tarsus, 39 ; bill 16. Colours : bill, black ; legs, black ; 
iris, dark brown. Stomach contents : small insects. 

The juvenile male is tawny black with the under-parts washed with honey- 
brown. Its primaries appear to be fully grown, but the central rectrices are absent. 

Crateroscelis robusta robust a (De Vis). 

Gerygone robusta De Vis, 1898, Ann. Rep. Brit. New Guinea, 1896-97, p. 84. Wharton Range, 
south-eastern New Guinea. 

1953. 17. 149-150 \ 2 $\ Mount Giluwe, 8,000-7,500 ft.; June, 1951. 

Measurements : 2 $ — wing, 64, 65 ; tail, 39, 40 ; bill, 12. Colours : bill, upper 
mandible dark horn and lower mandible light horn ; legs, flesh ; iris, yellow to 

Although these birds have been named robusta they differ from topotypical 
specimens by being more greyish in colour. In addition, the wing coverts, the lower 
part of the back, lower abdomen and flanks have a faint olivaceous wash. Mayr and 
Gilliard (1954 : 345) and Gyldenstolpe (1955 : 84) reported that among the birds from 
the Bismarck, Hagen and Kubor Mountains which they named robusta there was some 
variation in colour although a few specimens matched topotypical material. This 
pair from Mount Giluwe were sent to the Berlin Museum ; there Professor Stresemann 
kindly compared them with the type of albigula Reichenow from Schraderberg in the 
Sepik Mountains, which is now regarded as a synonym of robusta. He found that they 
agreed with this specimen so it might appear that albigula is a valid race, but in view 
of the variation in the series collected by Mr. Gilliard and Count Gyldenstolpe I 
consider that it is better not to separate albigula on these two specimens. 

Eupetes leucostictus loriae Salvadori 

Eupetes loriae Salvadori, 1896, Ann. Mus. Civ. Genova, (2) 16, (= 36), p. 102. Moroka, south- 
eastern New Guinea. 

1953. 17. 151, 153, 153a ; 1 (J, i §," 1 juv. ^ ; Mount Giluwe, 8,500 ft.; May, June> 


1953. 17. 152 ; 1 ? ; Hagen Range, 8,500 ft.; February, 1951. 

Measurements : 1 <J — wing, 76 ; tail, 82 ; bill, 18. 2 ? — wing, 84,86 ; tail, 94, 
98 ; bill, 18, 19. 1 juv. $ — wing, 82 ; tail, 84 ; bill, 17. Colours : bill, black ; 
legs, brown-grey to brown-black ; iris, dark brown. 

These birds are completing moult or are in fresh plumage except the February bird, 
where the plumage is worn. They were taken in the forests of the north slopes of 
Mount Giluwe. Nearest to loriae but differ by being a clearer blue below, due to the 
green wash being restricted to the flanks, the sides of the thorax and to a small patch 
on the upper breast near the throat. (The blue, however, is not so bright as that of a 
specimen of amabilis received on loan from the American Museum of Natural History.) 
The upper-parts appear to be a brighter green, slightly less olivaceous in colour ; 
this difference, however, may be due to fading on the part of the older specimens. 
The bills are somewhat more slender than in both topotypical loriae and amabilis, 
particularly the latter. 

Field note. Native name, " Penkeklempo " or "Kengnan" (the latter is 
probably the name for E. castanonotus) . A nest was found in a hole about 2 ft. above 
the ground in a half-dead stump of a tree-fern. An adult continued to visit the nest 
frequently and did not appear to be shy although the porters were nearby. It 
searched for insects on the ground by turning over moss and leaves with a quick 
movement of the bill ; insects that were taken were too small to be identified. It is 
possible that both parents feed the young because a second visit to the nest followed 
too soon upon the first for the same individual to have found food so quickly. When 
it returned to the nest the bird remained near the ground and jumped into the nest- 
hole from immediately beneath it. Faeces were removed from the nest on several 
occasions. The nest was situated so that no sunlight fell directly on it and the nest 
hole faced to the east. On 5th October some of the vegetation was removed to allow 
more light to fall on the nest and two young birds covered with a heavy grey down 
were discovered there. They crouched down in the middle of the nest at first, then the}^ 
stretched out their necks and started to hiss in a snake-like manner. The nest was 
then observed from a short distance away, after a while the juveniles started to preen; 
but when the sun began to fall directly on to the nest the young birds began to shift 
around and utter a harsh cry. Finally they jumped out of the nest and one of the 
parents lead them away along a small run through the undergrowth. The nest itself 
was circular in shape, with an external diameter of about 145 mm. and an internal 
one of about 75 mm.. It was heavily built on an untidy foundation of sticks, ferns and 
dead leaves, the inner cup was composed of fine rootlets lined with the skeletons of 
leaves, although one green leaf was removed from the nest. A second nest was found 
containing two young birds on 8th October ; these were about the same stage of 
development as the young in the first nest. It was situated in a nest-hole about 3 ft. 
from the ground and similarly built to the first one only with less foundation. Both of 
the parents fed the young. The gape of the juvenile was white and the inside of the 
mouth a pale orange. The young were fed eleven times in one hour at the height of 
feeding, but only five times between 11.50 and 12.50 hours. On one occasion one of 
the parents sat on the nest and started to brood the young, when the other came to 
feed the juveniles it pushed the brooding parent off the nest ; there appeared to be 


no recognition behaviour. The adults were very tame and continued their search for 
insects to within an arm's length of the hide and ignored all noises that came from 
inside. A third nest was found in a hole towards the base of a tree-fern. In it were 
two eggs (16th October), Mayr and Gilliard (1954 : 345). 

Ifrita kowaldi kowaldi (De Vis) 

Todopsis kowaldi De Vis, 1890, Ann. Rep. Brit. New Guinea, 1888-89, p. 59. Owen Stanley 
Mountains, south-eastern New Guinea. 

Field note. Native name, " Kompe-chicko ". A nest was found built about 12 
ft. up in a slender sapling, it was thickly decorated with moss which gave it a massive 
appearance. In the nest there was one juvenile which seemed to suffer badly from 
the heat. Both parents fed the young bird ; once the juvenile was given a cicada-like 
insect which was so large that the parent seemed to have difficulty in forcing it down 
the young bird's throat. 

Malurus alboscapulatus subspecies ? 

Malurus alboscapulatus A. B. Meyer, 1874, Sitzungsber. Akad. Wiss. Wien, 69, p. 469. Arfak 
Mountains, north-western New Guinea. 

1953. 17. 154-157 ; i cJ, 2 juv. (J, i ? ; Mount Giluwe, 7,500-7,300 ft.; April, June, 


Measurements : 3 <J — wing, 51-53 ; tail, 50-59 ; bill, n-5-i2-5. 1 ? — wing, 
49 ; tail, 57 ; bill, 11. Colours : bill, black (unsexed specimen, lower mandible 
dark horn) ; legs, dark horn ; iris, dark horn. Stomach contents : insects. 

Mayr and Gilliard (1954 : 346) and Gyldenstolpe (1955 : 88) reported that apart 
from wing size all the birds collected in the Wahgi region agreed with topotypical 
specimens of the race mafulu from the mountains of south-eastern New Guinea. It 
seems to me, however, that the adult male from Mount Giluwe is a closer match with 
a specimen of the longer winged race balim from the Snow Mountains which I had on 
loan from the American Musuem of Natural History than with a specimen of mafulu. 
But, again, the immature birds seem to be a fairly close match with young mafulu. 
The birds from Mount Giluwe are also larger than birds of the race tappenbecki (<£, 
wing, 45-48) from the lower Ramu and Sepik rivers north of the Schrader-Bismarck 
Range, an area considerably nearer than either the Snow Mountains or Mafulu. 
Professor Stresemann who kindly compared the series with topotypical specimens of 
tappenbecki in the Berlin Museum was able to separate the immature birds on colour 
as well as wing length. 

Megalurus timoriensis subspecies ? 

Megalurus timoriensis Wallace, 1863, Proc. Zool. Soc. Lond. 1863, p. 489. Dilly, Timor. 

Series A : 

1953. 17. 158 ; 1 (J ; Mount Giluwe, 12,000 ft.; April, 1951. 

1953 . 17 . 159-162 ; 1 juv. <J, 3 $ ; Lamende Range, 11,000 ft.; May, 1951. 


Series B : 

1953. 17. 163-166 ; 4 (J ; Mount Giluwe, 7,300 ft.; April, 1951. 

Measurements : Series A : 2 <$ — wing, 73, 74 ; tail, 95-5, 96 ; bill, 12, 14. 
3 ? — wing, 68-71 ; tail, 93-97 ; bill, 12. Series B : 4 <$ — wing, 71-75 ; tail, 
98-109 ; bill, 12-13. Colours : bill, upper mandible dark horn and the lower 
mandible pale horn ; legs, brownish dark flesh ; iris, Series A, light to dark brown — 
Series B, pale orange to light brown. Stomach contents : insects. 

Mayr and Gilliard (195 1 : 9) described two races of this species of warbler from 
Mount Hagen. They named the birds of the summit (alpine) grasslands at 12,000 ft. 
montanus and those of the grass slopes at 7,800 ft. wahgiensis. The birds were 
separated mainly on the somewhat darker, more rufous, colouring of the light areas 
of the upper-parts of montanus. Through the kindness of the American Museum of 
Natural History I have examined two specimens of montanus and one of wahgiensis 
from the series on which the descriptions were based. The specimens in Series A and 
Series B listed above are distinguishable from each other and, as far as I can see, the 
high altitude series matches Mayr and Gilliards' lower altitude form wahgiensis, 
while the lower series on Mount Giluwe matches the high altitude montanus. The 
explanation of this curious reversal on two neighbouring peaks is not evident. It 
seems certain, at least, that the differences are not caused by seasonal changes in 
plumage for in both series there are birds in worn and fresh dress. 

Field note. Native name, " Kuchaill " (" 11 " as in Welsh). 

Cisticola exilis diminuta Mathews. 

Cisticola exilis diminuta Mathews, 1922, Birds Austr. 9, p. 373. Paterson Creek, Cape York, 
Queensland, Australia. 

1953. 17. 167 ; i ? ; Mount Giluwe, 7,300 ft.; June, 1951. 

Measurements : 1 ? — wing, 44 ; tail, 47 ; tarsus, 18 ; bill, 9-5. Colours : 
bill, horn colour, darker on ridge ; legs, flesh colour ; iris, dull brown. Stomach 
contents : insects. 

Generally regarded as being a species restricted to the grasslands below 4,000 ft. 
it is interesting that this specimen was taken at 7,300 ft.. The specimen is in winter 

Sericornis perspicillatus Salvadori 

Sericornis perspicillata Salvadori, 1896, Ann. Mus. Civ. Genova. (2) 16 ( = 36), p. 99. Moroka, 
south-eastern New Guinea. 

Field note. A somewhat pear-shaped nest was found in the forest near Tomba. 
It was about 2 to 3 ft. above the ground, hanging by a few fibres from a twig. The 
entrance to the nest was about half-way up one side ; above it a small bulge formed 
a " porch ". Two young birds were in the nest (15th October). Both parents fed the 


young and during a period of one hour 16 visits were made with food ; nest sanitation 
occurring 3 times. During another period of watching for 30 minutes food was taken 
to the nest 7 times. Among the insects fed to the juveniles a moth, a small green 
caterpillar and a Tipulid were identified. The adults chattered noisily around the 
nest. They were active and seldom remained still, hopping around in the undergrowth 
and sideways up the stems of the bushes. They flirted their wings constantly, like 
birds of the genus Phylloscopus. 

Gerygone ruflcollis insperata De Vis 

Gerygone insperata De Vis, 1892, Ann. Rep. Brit. New Guinea, 1890-91, p. 94. Mount Suckling, 
south-eastern New Guinea. 

Field note. Native name, " W'un-monamp ". A domed nest containing two 
fully-fledged juveniles was found on 3rd October in the forest near an area of swamp- 
land. One point of interest is that three different adults appeared to feed the young 
birds. Apart from this the behaviour, of the adults was typical of most warblers ; 
they flitted with quick movements from branch to branch and never remained still 
for any length of time. Both the young and the adults were very noisy. 

Peltops montanus Stresemann 

Peltops blainvillii montanus Stresemann, 1921, Anz. Ornith. Ges. Bayern, 1, p. 35. Hunstein- 
spitze, Sepik Mountains, northern New Guinea. 

Field note. One bird was seen in the forest at the top of a high tree, the identity 
of this blackbird with a white patch below and behind each eye and the crimson 
rump and vent was unmistakable. 

Rhipidura brachyrhyncha devisi North 

Rhipidura devisi North, 1897, Proc. Linn. Soc. New South Wales, 22, p. 444. New name for 
Rhipidura albicauda De Vis, 1897, Ibis, p. 375. No type locality given (= ? Mt. Scratchley). 
(Not Rhipidura albicauda North, 1895). 

1953. 17. 168 ; 1 imm. <J ; Mount Giluwe, 7,500 ft.; June, 1951. 

Measurements : 1 imm. <$ — wing, 71-5 ; tail, 92 ; tarsus, 24 ; bill, 8. Colours : 
bill, upper mandible dark horn, lower mandible light horn ; legs, pale flesh ; iris, 
brown-black. Stomach contents : insects. 

Rhipidura albolimbata lorentzi van Oort 

Rhipidura albo-limbata lorentzi van Oort, 1909, Nova Guinea, 9, (Zool.), p. 85. Hellwig Moun- 
tains, Oranje Range, New Guinea. 

1953. 17. 169 ; i ? ; Mount Giluwe, 8,000 ft.; June, 1951. 

Measurements : 1 $ — wing, 82 ; tail, 85 ; bill, 10. Colours : bill, not recorded ; 
legs, black ; iris, brown-black. Stomach contents : insects. 


Field note. Native name, " Ipenem ". Two nests were found, one was built 
about 20 ft. above the ground in a tree, the other about 12 ft. up in a creeper. Each 
was a small neat cup with a " tail ", the outsides were close-knit in appearance as if 
spiders' webs had been woven in. Both nests contained young (26th September) 
which the adults continued to feed while the party stood beneath the nests. 

Rhipidura leucophrys melaleuca (Quoy and Gaimard) 
Muscipeta melaleuca Quoy and Gaimard, 1830, Voy. Astrolabe, Zool. 1, p. 180. New Ireland. 

1953. 17. 170 ; 1 $ ; Mount Giluwe, 7,300 ft.; June, 1951. 
Measurements : i $ — wing, 104 ; tail, 103 ; tarsus, 25 ; bill, 13. Colours : 

bill, black ; legs, black ; iris, brown-black. Stomach contents : insects. 

Usually regarded as a bird of the coast, lowland lakes, swamps and rivers it is 
interesting that Mr. Shaw-Mayer found it to be "fairly plentiful" on the high 
plateau near Mount Giluwe. Mayr and Gilliard (1954 : 348) reported that it was 
common at Nondugl at 5,000 ft., and that it was nesting at 7,000 ft. 

Field note. A common bird which spends most of the time on the ground. The 
tail is not spread out much compared with many other members of the genus. When 
the bird jumps up to a perch from the ground the long tail is swung pendulum- 
fashion in the manner of that of a shrike. 

Machaerirhynchus nigripectus saturatus Rothschild and Hartert 

Machaerirhynchus nigripectus saturatus Rothschild and Hartert, 1913, Novit. Zool. 20, p. 498. 
Mount Goliath, Snow Mountains, New Guinea. 

1953. 17. 171 ; i c? ; Mount Giluwe, 8,000 ft.; May, 1951. 

Measurements : 1 <J — wing, 64 ; tail, 62 ; bill, n. Colours : bill, black ; legs, 
black ; iris, black-brown. Stomach contents : small insects. 

Field note. Native name, " Koongnaremp ". Fairly common in the forest. 

Microeca papanua Meyer 

Microeca papuana A. B. Meyer, 1875, (April), Sitzungsber. Abh. Nat. Ges. Isis, Dresden, 1875, 
p. 74. Arfak Mountains, north-western New Guinea. 

I 953- I 7-i72 ; i(J; Mount Giluwe, 7,500 ft.; May, 1951. 

Measurements : 1 <J- — wing, 78-5 ; tail, 48 ; bill, 10. Colours : bill, black ; 
legs, yellow ; iris, dark brown. Stomach contents : insects. 

This specimen is a little larger and darker in colour than the other birds of this 
species that I have seen and there is marked reduction in yellow on the outer webs 
of the primaries. Rand (1942& : 483) reported similar differences in the birds he 



collected in the mountains of Dutch New Guinea and correlated the increase in size 
with altitude. He suggested that his birds appeared to be darker because the material 
with which they were being compared was foxed, and this may well account for the 
colour difference between the birds before me. 

Petroica bivittata bivittata De Vis. 
Petroeca bivittata De Vis, 1897, Ibis, p. 376. Mount Scratchley, south-eastern New Guinea. 

1953. 17. 173, 176 ; 1 (J, 1 ? ; Mount Giluwe, 11,000-10,000 ft.; April, May, 1951. 

1953. 17. 174-175 ; 2 (J ; Lamende Range, 11,000 ft.; May, 1951. 

Measurements : 3 <J — wing, 73-77 ; tail, 46-48 ; bill, 9-0. 1 ? — wing, 73 ; tail, 
45 ; bill, 9-0. Colours : bill, black ; legs, black ; iris, dark brown. Stomach 
contents : small insects. 

The April bird (sex ?) has started moulting, one bird taken in May is in fresh 
plumage the other two are completing moult. Collected on the alpine grasslands. 

Peneothello sigillatus subspecies ? 

Poecilodryas (?) sigillatus De Vis, 1890, Ann. Rep. Brit. New Guinea, 1888-89, p. 59. Mount 
Victoria, south-eastern New Guinea. 

Field note. An adult female was seen on 18th October on an empty nest which 
was half concealed in moss and creeper. The nest was built of moss, green ferns and 
rootlets, with a thick lining of moss and dead fern leaves. It measured approximately 
50 mm. in diameter internally and 115 mm. externally, the cup being about 25 mm. 

Peneothello cyanus subcyaneus (De Vis) 

Poecilodryas subcyanea De Vis, 1897, Ibis, Ser. 7, 3, p. 377. Mountains of south-eastern New 

1953. 17. 177 ; i <J ; Mount Giluwe, 7,500 ft.; May, 1951. 

Measurements : 1 $ — wing, 99 ; tail, 68 ; bill, 15. Colours : bill, black ; 
legs, black ; iris, brown-black. Stomach contents : small insects. 

Field note. Native name, " Goorgla ". Nests were built in the forks of upright 
branches in the bushes, and appeared to be made mainly of fine rootlets covered with 
moss and lined with dried fern fronds. They were round and deep, being about 55 
mm. in diameter and 45 mm. deep internally and 130 mm. in diameter and 80 mm. 
high externally. One nest, first examined on 18th September, contained one juvenile 
almost ready to fly ; it was visited daily until 22nd September, when the young 
bird had flown. Another nest had one juvenile in it on 28th September, but this bird 


had flown when the nest was examined on the following day. An egg was found in a 
nest 17th October. It was a pointed oval with a light olive ground colour ; there 
Mere a few scattered dark brown spots and a fainter red-brown speckling that became 
more intense towards the broader end. 

Heteromyias albispecularis centralis Rand. 

Heteromyias albispecularis centralis Rand, 1940, Amer. Mus. Novit. No. 1074, p. 4. Near 
Bernhard Camp, Idenburg River, at 2,150 m., central New Guinea. 

1953. 17. 178-180 ; i o, i juv. cJ, i $ ; Mount Giluwe, 8,000-7,800 ft.; April, 
June, 1951. 

Measurements : 2 <J< — wing, 99, 101 ; tail, 61, 62 ; bill, 17, 18. 1 ? — wing, 98 ; 
tail, 58 ; bill, 17. Colours : bill, black with tip pale horn (juv. <J, pale horn) ; 
legs, flesh ; iris, dark brown. Stomach contents : small beetles and other insects. 

The two April birds are completing moult, although a few juvenile feathers still 
remain in the young male. The adult male (June) is in fresh plumage. 

Field note. Native name " Yamari." Several nests were found in dark parts of 
the moss forest where the light-meter read as low as 3-2 foot-candles for the most of 
one forenoon. They were built 7 to 10 ft. above the ground in the upper branches of 
young saplings, although one was found in a small bush where it was only 3 ft. from 
the ground. The nests appeared to be very similar in being heavily built on slender 
branches, which in some cases could not adequately support their weight. One nest 
that was examined was found to be about 90 mm. in diameter and 40 mm. deep ; 
it was built of grass and twigs with a little moss and lined with thin fern rootlets. All 
the nests contained young birds. One juvenile examined on 20th September was 
blackish brown above with the shafts of the feathers lighter in colour. The breast 
was brown and the belly white with a brown wash. The gape and the inside of the 
mouth were yellow. The legs, which were long and strong, were flesh coloured. 
The irides were brown. Another nest, seen two days later on 22nd September, 
contained only one nestling which was quite naked ; yet a week later, on 29th 
September, a nest was found with one juvenile which had well-grown wing quills. 
This young bird rose on its legs and with arched wings started to cheep when the 
parents returned to the nest. It was established that both parents fed the young 
whose main diet appeared to consist of earthworms. On two occasions a juvenile was 
seen to eject a bag of faeces on to the rim of the nest, one of the adults returned to 
the nest expressly to remove it — or so it appeared. The adults were never seen in the 
forest except at the nest. Sometimes they would return to the nest silently, at other 
times they would call quietly with a soft, musical piping sound. The young always 
answered this call, but not loudly. When the adult made this piping call the throat 
vibrated, but the bill remained closed. The alarm call was, " kak-kaktak-kak ". 
Once when a nest was being examined an adult repeatedly exhibited a distraction 
display by " injury feigning " on the ground where it dragged one wing as if it were 
zool. 3, 10. 27 


The adults had a habit of remaining hidden in the undergrowth and of standing 
quietly on the nests, a form of behaviour that is more characteristic of a babbler than 
a flycatcher. The bird's long strong legs and heavy bill are other characters that 
suggest that this bird may have some affinity with the Timalinae. As opposed to 
these Timaloid tendencies adults were seen to perch sideways on the moss-covered 
trunks of the trees in a position curiously like that of a nuthatch. 

Pachycephala schlegelii obscurior Hartert 

Pachycephala schlegelii obscurior Hartert, 1896, Novit. Zool. 3, p. 15. Eafa District, Owen 
Stanley Mountains, south-eastern New Guinea. 

1953. 17. 181 ; i (J ; Mount Giluwe, 7,500 ft.; May, 1951. 

Measurements : 1 <J — wing, 88 ; tail, 65 ; bill, 11. Colours : bill, black ; 
legs, brown-black ; iris, brown-black. Stomach contents : insects. 

Pachycephala modest a hypoleuca Reichenow 

Pachycephala hypoleuca Reichenow, 1915, Journ. f. Ornith. 63, p. 125. Schraderberg, Sepik 
Mountains, northern New Guinea. 

1953. 17. 182-183 ; i (J, i ? ; Lamende Range, 11,000 ft.; May, 195 1. 

Measurements : 1 <J — wing, 90 ; tail, 64 ; bill, n. 1 ? — wing, 89 ; tail, 65 ; 
bill, 12. Colours: bill, black; legs, dark grey to black ; iris, dark brown. Stomach 
contents : small grubs and insects. 

These specimens were collected among the small trees on the alpine grasslands. 

Pachycephala rufinucha niveifrons Hartert 

Pachycephala rufinucha niveifrons Hartert, 1930, Novit. Zool. 36, p. 57. Wondiwoi, Wandam- 
men Mountains, north-western New Guinea. 

1953.17. 184 ; 1 juv. <J ; Hagen Range, 8,000 ft.; March, 1951. 

1953. 17. 185-186 ; 2 (J ; Mount Giluwe, 8,000-7,8000 ft.; April, 1951. 

Measurements : 3 <J — wing, 89-91 ; tail, 68-70 ; bill, 16-5, (juv., 14). Colours : 
bill, black (juv., horn) ; legs, olivaceous grey-black (juv., horn) ; iris, yellow (juv., 
dark grey). Stomach contents : insects. 

Apart from having slightly longer wings these birds match material taken at lower 
altitudes on the Utakwa River (wing, 84-86). Rand (19426 : 489) showed that in this 
species wing length could be correlated with altitude, the birds from the higher 
altitudes being slightly larger. 

Field note. A nest was built in a fork of a branch about 7 ft. up in a bush. The 
outer part of the nest was made of ferns, coarse rootlets and green moss, the inner 
part of the thin fibres, fine rootlets and the skeletons of leaves. There were two well 



incubated eggs in the nest on 17th October. These were a chalky white colour with 
black spots and a fainter underlay of grey spots. Another nest found two days later, 
on 19th October, contained two juveniles ; the insides of their mouths were a pale 
orange apart from two prominent blue spots at the posterior of the palate. It is 
probable that the young were fed by both parents, but the nest was visited only 
infrequently. During a three and a half hour watch on the nest an adult visited it 
only four times during the first half hour, three times during the second, once during 
the third but four times during the last half-an-hour. 

Myiolestes megarhynchus tappenbecki (Reichenow) 

Colluricincla tappenbecki Reichenow, 1899, Journ. f. Ornith. 47, p. 118. Friedrich Wilhelms 
Hafen (= Madang), Astrolabe Bay, north-eastern New Guinea. 

1953. 17. 187-190 ; 2 cJ, 2 ? ; Minj, Wahgi River, 5,000 ft.; November, 1950. 

1953. 17. 191 ; 1 ? ; Mount Giluwe, 7,5000 ft.; May, 1951. 

Measurements : 2 & — wing, 94, 96 ; tail, 76, 79 ; bill, 20, 21. 3 ? — wing, 91 — 
92 ; tail, 72 — 76 ; bill, 18 — 19. Colours : bill, 2 £, brown horn — 3 $, horn ; legs, 
blue-grey to dark grey ; iris, brown. Stomach contents : insects ; 1 $, small 
(?) water snail. 

All the four birds taken at Minj in November, 1950, were taken on the nest, the 
nests being built about one foot above the ground in long grass. It is interesting to 
note that these four birds are in different plumages. The two males are in full moult ; 
and of the two November females one is in worn dress while the other is in fresh 
plumage. The plumage of the female taken in May shows some signs of wear. 

Eulacestoma nigropectus subspecies ? 

Eulacestoma nigropectus De Vis, 1894, Ann. Rep. Brit. New Guinea, 1893-94, p. 102. Mount 
Maneao, south-eastern New Guinea. 

1953. 17. 192 ; ? ? ; Mount Giluwe, 8,000 ft.; May, 1951. 

Measurements : 1 ? ? — wing, 71 ; tail, 52 ; bill, 11-5. Colours : bill, horn ; 
legs, purple-flesh ; iris, grey-brown. Stomach contents : insects. 

This is a juvenile, and in the absence of comparable material I cannot name it 
racially. It is probably clara Stresemann and Paludan as Mayr and Gilliard (1954 : 
354) reported that their birds from this area were that race. 

Lanius schach stresemanni Mertens 

Lanius schach stresemanni Mertens, 1923, Senckenbergiana, 5, p. 228. Kulungtufu, Saruwaged 
Mountains, north-eastern New Guinea. 

1953. 17. 193, 195-196 ; 1 <J, 2 ? ; Mount Giluwe, 7,300 ft.; May, June, 1951. 
1953. 17. 194 ; 1 juv. cJ ; Wahgi River, 5,200 ft.; November, 1950. 

ZOOL. 3, IO . 27§ 


Measurements : i $ — wing, 101 ; tail, 140 ; bill, 16. 1 juv. <J — wing, 95 ; 
tail, 105 ; bill, 13. 2 $ — wing, 97, 98 ; tail, 121, 123 ; bill, 15. Colours : bill, 
black (juv. (J, lower mandible, horn) ; legs, black ; iris, brown-black. Stomach 
contents : small grubs, beetles and other insects. 

" Not many of this solitary species on the grasslands of the plateau. A bird here 
and there, as usual clinging to the stem of cane-grass." (F. S.-M.). The adult male 
is in fresh plumage and the testes are slightly enlarged. The plumage of the young 
male is worn. The two females are completing moult with the outer primaries 

Field note. Native name, " Komkompir ". Two deep-cupped nests, built about 
6 ft. above the ground, were found in tall reeds. One, examined on 5th September, 
contained two eggs; but in the other, found on 23rd September in pit-pit country, 
were two well-grown juveniles. 

Artamus maximus Meyer 

Artamus maximus A. B. Meyer, 1874, Sitzungsber. Akad. Wiss. Wien, 69, p. 203. Hatam, 
Arfak Mountains, north-western New Guinea. 

1953. 17. 197 ; i c? ; Minj, Wahgi River, 5,000 ft.; November, 1950. 

Measurements : 1 (J- — wing, 164 ; tail, 73 ; bill, 23. Colours : bill, blue-grey ; 
legs, grey ; iris, dark brown. Stomach contents : small beetles. 

Gyldenstolpe (1955 : 121) separated A. m. wahgiensis mainly on the darker colour 
of the upper-parts of the birds from the Wahgi region in comparison with a series from 
the Arfak Mountains. The present specimen, however, hardly differs from the Arfak 
Mountain bird. The only variations in colour in the specimens I have examined 
appear to be correlated with age and season. In a series from south-eastern New 
Guinea there are some individuals lighter in colour than both the Wahgi River and 
Arfak birds, and others that are darker in colour. It seems that the lighter coloured 
birds are juveniles or adults in worn, faded plumage ; while the darker birds are 
adults in fresh plumage. Unfortunately this series is incomplete in specimens taken 
in all months of the year, so the degree of seasonal variation cannot be established 
with certainty. As the individual variation is greater than that between the birds 
from the Central Highlands and the Arfak Mountains, I am not naming the present 
bird racially. 

Field note. Native name, " Kompurkaikir ". A nest was built about 50 ft. 
above the ground in the fork at the base of an upright branch of a dead tree in the 
forest. Three juveniles, almost ready to fly, were found in it (26th September). It is 
interesting that four or five different adults were observed to feed the young, and on 
one occasion three adults were seen on the nest at the same time ! Communal nesting 
of this nature is rare and provides a subject for further investigation. Like many 
other communities this one was very noisy, with both the adults and the juveniles 
contributing to the chatter. The juvenile plumage was similar to that of the adult, 
but the dark areas of the body were necked with a light brown ; the bill of the young 
bird was horn coloured. 


Aplonis metallica metallica (Temminck) 

Lamprotornis metallicus Temminck, 1824, Planch. Col. d'Ois, pi. 266. " Celebes," by error for 
Amboina, Moluccas. 

1953. 17.198 ; i ? ; Mount Giluwe, 7,000 ft.; May, 1951. 

Measurements : 1 ? — wing, 104 ; tail, 91 ; bill, 16. Colours : bill, black ; 
legs, black ; iris, bright orange-red. 

" Few Aplonis were seen at 7,000 feet when this specimen was taken in the wood- 
lands but the bird was plentiful at 5,300 feet." (F. S.-M.). 

Pomareopsis briujni (Salvadori) 

Grallina bruijni Salvadori, 1875, Ann. Mus. Civ. Genova, 7, p. 929. Arfak Mountains, north- 
western New Guinea. 

Field note. Native name, " Notechll " (" 11 " as in Welsh). This is a bird of the 
mountain stream, and one was seen by a rushing torrent at about 6,500 ft. When it 
alighted on a rock it fanned out its tail in the same manner as the Plumbeous Red- 
start, Pheonicurus fuliginosus , of the fast-flowing streams of the Himalayas. 

Epimachus meyeri bloodi Mayr and Gilliard 

Epimachus meyeri bloodi Mayr and Gilliard, 1951, Amer. Mus. Novit. No. 1524, p. 10. Mount 
Hagen, Central Highlands, 8,300 ft., New Guinea. 

1953. 17. 199-200 ; i (J, i juv. (J ; Hagen Range, 9,000 ft.; February, 1951. 

1953. 17.201 ; 1 ? ; Mount Wilhelm, Bismarck Range, 8,000 ft.; May, 1951. 

Measurements : 1 # — wing, 174 ; tail, 604 ; bill, 78. 1 juv.<J — wing, 149 ; 
tail, 320 ; bill, 60. 1 °- — wing, 149 ; tail, 312 ; bill, 76. Colours : bill, black ; legs, 
dark grey (ad. <$ black) ; iris, blue-grey. Stomach contents : fruits. 

The juvenile male resembles a series of adult females from the Hagen Range except 
in the length of the bill. The adult female from the Bismarck Range differs from this 
series in the upper-parts being lighter, less olivaceous, more like the ochraceous 
colour of the females of meyeri. Not until more material is available from the Bis- 
marck Range will it be possible to assess the importance of this difference, but it may 
be that the Wahgi Gap country plays just as an important part in the distribution of 
the races of E. meyeri as it does in the distribution of the races of Astrapia stephaniae, 
(see below). The plumage of the present birds is fairly fresh, showing little signs of 
wear. Specimens taken in October in previous collections from Mount Hagen are in 
worn plumage and those taken in November are in full moult. It appears from the 
stomach contents of the birds that E. meyeri is not entirely frugivorous, for the 
stomachs of adult birds have been found to contain berries, grasshoppers and other 
insects in addition to various fruits. 


Astrapia stephaniae femina Neumann 

Astrapia stephaniae femina Neumann, 1922, Verh. Ornith. Ges. Bayern, 15, p. 236. Schrader- 
berg, Sepik Mountains, 60 miles north of the Hagen Mountains, northern New Guinea. 

1953.17.202 ; 1 $ ; Mount Wilhelm, 8,000 ft.; May, 1950. 

Measurements : 1 ? — wing, 145 ; tail, 320 ; bill, 24. Colours : bill, black ; 
legs, blue-grey (feet, black) ; iris, black-brown. Stomach contents : fruits. 

Mayr and Gilliard (1952a) showed that the grasslands between the Hagen Range 
and the mountains of the Wahgi Divide (a westerly spur of the Bismarck Range) 
acted as a barrier in the distribution of A. mayeri, a species closely related to A. 
stephaniae. The same authors (1954 : 321) also described the deforestation in the 
Wahgi Valley, which they suggested formed another important barrier to forest 
birds. It appears, then, that the mountain forest birds of the Kubor Range along the 
south and the Hagen Range at the head of the Wahgi Valley are isolated by open 
country from the mountain forest birds of the Wahgi Divide and the Bismarck Range 
north of the valley. This barrier apparently influences the distribution of the races 
of A. stephaniae, for specimens collected in the Bismarck Range by Mr. F. Shaw- 
Mayer in 1946 have been matched with cotypical material of femina, and Gylden- 
stolpe (1955 : 127) identified his birds from north of the Wahgi Valley as this race. 
On the other hand, two specimens from Mount Giluwe (see below) appear to be 
ducalis, which is the race found to the south of the valley in the Kubor Range (Mayr 
and Gilliard, 1954 : 356). The adult females from the Bismarck Range are slightly 
more olivaceous above than the adult females of ducalis and the under-parts are rust 
coloured barred with black instead of a buffy colour barred with black. In the colour 
of the under-parts they agree with the differential characters given by Mayr and 
Gilliard (1954 : 356). (They do not, however, exhibit the subobsolete white streaks 
at the bases of the central tail feathers described by these authors.) The present 
specimen differs somewhat from other material in having a narrow chestnut collar 
barred with black across the shoulders, the colour of the collar becoming lighter 
laterally as it merges into the rust barred-black of the breast. 

Astrapia stephaniae ducalis Mayr 

Astrapia stephaniae ducalis Mayr, 1931, Mitt. Zool. Mus. Berlin, 17, p. 711. Dawong, Herzog 
Mountains, eastern New Guinea. 

1953.17.203-204 ; i (J, i $ ; Mount Giluwe, 8,500 ft.; March, April, 1951. 

Measurements : 1 $ — wing, 165 ; tail, 660 ; bill, 20. 1 $ — wing, 137 ; tail, 
320 ; bill, 21. Colours : bill, black ; legs, dark grey ; iris, brown-black. Stomach 
contents : fruits and berries. 

Field note. Five birds, females or immature males, were seen in the forest. In 
flight, with their long tails streaming behind, they were reminiscent of the Racquet- 
tailed Drongo, Dissemurus paradiseus ; the diving flight being particularly like that 


of the Dicruridae. The birds appeared to be hunting for insects in the moss covering 
the branches of the trees ; one bird was seen to swallow what was probably a spider. 

Astrapia mayeri Stonor 

Astrapia mayeri Stonor, 1939, Bull. Brit. Ornith. Club, 59, p. 57. " Eighty to a hundred miles 
west of Mount Hagen," central New Guinea. 

1953.17.205 ; 1 c? ; Mount Giluwe, 9,000 ft.; March, 1951. 

1953.17.206-207 ; 1 juv. (J, 1 ? ; Hagen Range, 9,000-8,500 ft.; December, 1950. 

Measurements : 1 cJ — wing, 177 ; tail, 852 ; bill, 13. 1 juv. <$ — wing, 170 ; tail, 
350 ; bill, 12. 1 ? — wing, 148 ; tail, 310 ; bill, 15. Colours : bill, black ; legs, 
blue-grey (feet, black) ; iris, brown-black. Stomach contents : fruits and green 

Field note. The branches of the forest trees on which the bird perches are 
thickly covered with moss. It is probably this layer of moss that prevents the long 
tail feathers from becoming damaged. 

Astrapia stephaniae x Astrapia mayeri 

1953.17.208 ; 1 (J ; Hagen Range, 8,000 ft.; March, 1951. 

Measurements : 1 $ — wing, 167 ; tail, 821 ; bill, 18. Colours : bill, black ; 
legs, dark grey (feet black) ; iris, brown-black. 

Close to mayeri in having the tail white for most of its length ; but the rachis of 
the distal two-thirds of the central rectrices are brown in colour, and the webs of 
these feathers are fringed irregularly with brown. The nasal tuft is reduced in size 
to that of an immature male of mayeri, that is intermediate between the condition in 
meyeri where the tuft is present and in stephaniae where it is absent. The primaries 
are darker than in mayeri, being a solid black with a purple lustre like those of 
stephaniae. Too much importance should not be attached to the proportion of hybrid 
specimens in collections, for in the field they occur in a limited area which happens 
to be one of the most accessible. Mr. Shaw-Mayer wrote (in correspondence 5th 
November, 1952) of the difficulties he had experienced in obtaining a series of hybrids, 
and that, " Over the greater part of their extensive range the species remain distinct. 
The overlapping areas where hybridism takes place are small. ... ". 

Paradisaea apoda salvadorii Mayr and Rand 

Paradisaea apoda salvadorii Mayr and Rand, 1935, Amer. Mus. Novit. No. 814, p. 11. Vanumai, 
Central Division Papua, south-eastern New Guinea. 

1953.17.209. i $ Wahgi River, 5,500 ft.; November, 1950. 
Measurements : 1 ? — wing, 161 ; tail, 118 ; bill, 32. Colours : bill, blue-grey ; 
legs, brown-horn ; iris, bright yellow. Stomach contents : fruits. 


" Count Raggi's Bird-of-Paradise was fairly plentiful on the Wahgi River plateau ; 
finding sanctuary in the casuarina groves, small stands of secondary forest and in the 
few remnants left of the original oak forests. By mid-November (1950) they were 
breeding and several nests were observed ; in nearly every case the nests were high 
up, 50 to 60 feet, in casuarina trees. Each nest contained only one egg." (F. S.-M.). 

Pteridophora alberti hallstromi Mayr and Gilliard 

Pteridophora alberti hallstromi Mayr and Gilliard, 1951, Amer. Mus. Novit. No. 1524, p. 12. 
Near Tomba, Hagen Range, Central Highlands, New Guinea. 

1953. 17. 210 ; i juv. (J ; Mount Wilhelm, 7,500 ft.; May, 1950. 

1953. 17. 211 ; 1 ? ; Hagen Range, 9,000 ft.; February, 1951. 
Measurements : 1 juv. <? — wing, 112 ; tail, 80 ; bill, 15. 1 $ — wing, 122 ; tail, 

81 ; bill, 15. Colours : bill, black ; legs, brown-black (juv. $, grey-brown) iris, 
dark brown ; inside of mouth of $, green. Stomach contents : green berries. 

Field note. Native name, " Kongbuk " (adult male); " Doongbi " (adult 
female and juveniles). An adult male was seen in the forest at about 7,000 ft. The 
bird was preening and called occasionally, the sound was like the squeaking of rusty 
iron. The long head plumes were manoevured to avoid the branches of the tree. A 
solitary bird. 

Loria loriae amethystina Stresemann 

Loria loriae amethystina Stresemann, 1934, Ornith. Monatsber. 42, p. 144. Schraderburg, Sepik 
Mountains, northern New Guinea. 

1953. 17.212 ; 1 (?; Mount Wilhelm, 6,500 ft.; May, 1950. 

1953. 17. 213 ; 1 cJ ; Hagen Range, 9,000 ft.; February, 1951. 
Measurements : 2 <J -wing, 103, 106 ; tail, 71, 72 ; bill, 15-5, 16. Colours : 

bill, black ; mouth- wattles, creamy white ; legs, dark olive ; iris, dark brown. 
Stomach contents : berries. 

Field note. A domed nest with an exceptionally wide round opening was built 
about 12 ft. up in a decayed tree-stump in the forest. It was deep-set in living moss 
and built of green ferns and moss with a lining of fern stalks. There was one juvenile 
in the nest (25th September). Its head and breast were a smoke-grey and the wings a 
darker grey ; the gape and the inside of the mouth were white in colour. While the 
nest was being watched during the early part of one morning the female returned 
about every 10 to 15 minutes to feed the young and called with a soft " wark, wark " 
each time she approached. The food was regurgitated and appeared to consist of 
large berries and plum-coloured fruits. A short while after being fed the young bird 
expelled the fruit stones from its crop, they were ejected with a force sufficient to 
shoot them out through the mouth of the nest. Faecal capsules were also ejected 
occasionally from the nest, but generally the female assisted with nest sanitation. 
When the nest was examined a few days later it was deserted and contained faeces and 
fruit stones. 


Cnemophilus macgregorii kuboriensis Mayr and Gilliard 

Cnemophilus macgregorii kuboriensis Mayr and Gilliard, 1954, Bull. Amer. Mus. Nat. Hist. 103, 
p. 361. Mount Orata, Kubor Mountains, Central Highlands, New Guinea, + 9,000 ft. 

1953. 17.214 ; 1 (J ; Minj River, Kubor Range ; 8,000-9,000 ft.; October, 1950. 
Measurements : i $ — wing, 109 ; tail, 88 ; bill, 13 ; crest, 37. Colours : bill, 

black ; legs, dark olive ; iris, dark brown. Stomach contents : fruits. 

Mayr and Gilliard separated this race on the paler colour of the upper-parts in 
comparison with sanguineus from Mount Hagen. In colour it is closer to the scarlet- 
vermilion of sanguineus than the pale orange of macgregorii from south-eastern New 
Guinea ; the colour of kuboriensis being orange- vermilion. This specimen was taken 
near the headwaters of the Minj River and it matches a series from the Bismarck 
Range, which suggests that there may be an east to west cline from the pale colour of 
macgregorii in the south-east to the deep, rich colour of sanguineus from the Hagen 
Range. The series consists of birds in new and worn plumage but there is little 
seasonal colour difference. 

Cnemophilus macgregorii sanguineus Iredale 

Cnemophilus macgregorii sanguineus Iredale, 1948, Australian Zool. 11, p. 162. Kumdi, Mount 
Hagen district, Central Highlands, New Guinea. 

1953. 17.215 ; i juv. <J ; Hagen Range, 9,000 ft.; February, 1951. 

1953. 17.216 ; 1 imm. <J ; Mount Giluwe, 9,000 ft.; May, 1951. 
Measurements : 1 imm. $ — wing, 109 ; tail, 85 ; bill, 20 ; crest, 36. 1 juv. 

cJ- — wing, 113 ; tail, 87 ; bill, 20 ; crest, 21. Colours : bill, black (juv. (J, tip 
dark horn) ; legs, brown-black to glossy black ; iris, dark brown (juv. <J, dull grey). 
Stomach contents : fruits. 

The young bird is a juvenile and indistinguishable from an adult female. The 
immature bird could well be described as " sub-adult ", for in colour it is intermediate 
between the young bird and an adult male, but the collector has indicated on the 
field label that the testes were as large as those of a breeding adult male. From this it 
seems that the bird comes into breeding condition before it is fully adult. If this is 
so it would be interesting to determine its behaviour towards an adult female and 
her response. The primaries and rectrices of this specimen are as richly coloured as 
topotypical sanguineus ; that is, darker than kuboriensis where these feathers are 
nearer the colour of macgregorii. It is on this character that I have been able to 
determine the racial identity of the bird. 

Archboldia papuensis sanfordi Mayr and Gilliard 

Archboldia papuensis sanfordi Mayr and Gilliard, 1950, Amer. Mus. Novit., No. 1473, p. 1. Four 
miles west of Tomba, south-western slope of Mount Hagen, Central Highlands, New Guinea. 

1953. 17.217, 218, 220, 222; 3 <J, 1 imm. <J ; Mount Giluwe, 9,000-8,500 ft.; 
March, April, May, 195 1. 


1953. 17. 219, 221, 223, ; i (J, i juv. (J, i imm. $ ; Hagen Range, 9,000-8,800 feet ; 
April, 1951. 

Measurements : 4 <J — wing, 165-171 ; tail, 167-171 ; bill, 17-20. 1 juv. 
(J — wing, 142 ; tail, 131 ; bill, 19. 1 imm. $ — wing, 163 ; tail, 141 ; bill, 22. 1 imm. 
$ — wing, 154 ; tail, 132 ; bill, 19. Colours : bill, black ; legs, grey-blue (imm. ?, 
bright light blue) ; iris, dark brown. Stomach contents : fruits. 

These are the first specimens of this genus to be received by the British Museum 
(Natural History). In comparison with the description of this race the feathers of 
the crest in the male exhibit some variation. Although they are predominantly 
black-based with orange-gold tips some have an area of white between the black and 
the orange-gold and a few are tipped with black. All the specimens are in moult. 

Field note.- — Near Tomba an area of the forest floor had been cleared to form a 
bower measuring approximately 5 ft. by 4 ft. The branches of the surrounding trees 
and a fallen tree trunk were densely decorated with the bamboo-like trailing vines of 
epiphytes which grow on high trees. Several branches were quite bare and smooth, 
probably from wear when the birds alighted and postured. The floor of the bower 
was strewn with the dead fronds of ferns. A number of shells of land snails were also 
found on the ground. 

Amblyornis macgregoriae De Vis 

(Text-fig 2) 

Amblyornis macgregoriae De Vis, 1890 (22nd February), Ann. Rep. Brit. New Guinea, 1888-89, 
p. 61. Musgrave Range, south-eastern New Guinea. 

1953.17.224 ; 1 cJ ; Rubor Range, 7,000 ft.; November, 1950. 

1953.17.225-226 ; i<$, 1 juv.cJ; Hagen Range, 9,000 ft.; February, 1951. 

1953.17.227 1 (J ; Lamende Range, 9,000 ft.; June, 1951. 

Measurements : 4 <J — wing, 134-137 ; tail, 83-86 ; bill, 17-19 ; crest, 60-65. 
Colours : bill, dark horn lighter towards base ; legs, dark olive-brown ; iris, dark 
brown. Stomach contents : fruits. 

Mayr and Gilliard (1954 : 363) discussed the small variations in colour and measure- 
ment found in this species. They considered that these were of a clinal nature and 
concluded that because of this it would be inadvisable to distinguish their birds from 
the Central Highlands as a separate race, although they differed somewhat from other 
populations. The material I have examined supports this conclusion, the underparts 
of the birds from the Central Highlands being, on the whole, rather more ashy in 
tone when compared with other populations. This difference however, is not constant 
and there is some individual variation for two adult males from Tomba match birds 
from the south-east. Although I have few adult males for comparison their measure- 
ments do not differ significantly from those of the birds from the Central Highlands. 

Measurements of adult males of Amblyornis macgregoriae 

Wing. Tail. Crest. 

macgregoriae Owen Stanley Range (2) . . 132, 140 . 80 . 57, 61 

aedificans Upper Waria River (2) , , 132, 138 . 80, 81 . 60, 61 



^•>&-jv>V/:.>: : :-:;y:i : :-::^<:i\^ 

2^i 260 .4-»--230 — *c 


Fig. 2. — Sketch of a cross-section of the bower of Amblyornis macgregoriae. 
Measurements in millimetres. (After Loke Wan Tho.) 

Field note. Bowers were found on several coccasions ; they were roughly 
circular in plan, but the dancing ground invariably formed a perfect circle. Each 
bower was built around a central sapling of Garcenia which the natives call ' ' Kichan' ' . 
One bower found deep in the moss forest was built of a dark moss and the playground 
was devoid of leaves. 

Chlamydera lauterhachi lauterbachi Reichenow 

Chlamydera lauterbachi Reichenow, 1897, Ornith. Monatsber. 
Ramu River, Central Highlands, New Guinea. 

14. Jagei River, upper 

1953.17.228-230 ; i (J, 2 ? ; Minj, Wahgi River, 5,000 ft.; November, 195 1. 

Measurements : 1 £ — wing, 133 ; tail, 104 ; bill, 22. 2 ? — wing, 126, 129 ; 
tail, 102, 103-5 J bill, 22. Colours : bill, black ; legs, olive-green ; iris, brown. 
Stomach contents : fruits and berries. 

" This bower-bird is found in lightly wooded country. It was plentiful on the 
plateau drained by the Wahgi River, where it frequented small patches of secondary